Insects under global warming. Review and retrospective analysis

We have tried to assess the morphological changes in insects under conditions of global warming using the example of three mass species. We analyzed collection funds for 140 years from the same territory in Western Siberia. We found out that the climate of Siberia is systematically drying up and the average temperatures are increasing, and as a result, the species composition of the fauna of Western Siberia changes, as well as the morphology of some insect species.


Introduction
Insects are highly adaptable to changing environmental conditions. Knowing the reactions of insects to changes in a particular factor, it is possible to predict or describe changes in the biocenosis. Biologists around the world are now studying the effects of climate change extensively. Entomologists have long known the morphological reaction of the honey bee Apis mellifera Linnaeus, 1758 to an increased temperature in the process of ontogenesis -the venation of the wing changes into the flesh until the appearance of new veins and cells (the exact direction of the process and the final result depending on temperatures is currently being studied). The study of the process of faunogenesis (tracking the species of invaders) can also serve to describe the consequences of climate change, along with changes in the physiology of a number of insect species. We tried to determine what morphological changes a species undergoes within one geographic point for 120 years under conditions of climate transformation, and also to review changes in the abundance of some species and in the fauna of Western Siberia.

Materials and Methods
To describe morphological changes, 3 cosmopolitan species (one species from three families) with a high population density were chosen as the object of research: Pompilidae -Anoplius (Arachnophroctonus) viaticus (Linnaeus, 1758); Sphecidae -Ammophila sabulosa (Linnaeus, 1758); Vespidae -Vespa crabro Linnaeus, 1758. For each species, three groups of individuals were analyzed: group no. 1 -wasps collected in 1893, group no. 2 -wasps collected in 1956, group no. 3 -wasps collected in 2017 (Fig. 1). The minimum number of specimens in the sample was 20 (A. sabulosa, group of 1893), the maximum -60 (V. rabro, group of 1893) (Table 1). For each individual, morphometric measurements of the fore wing (right and left) were carried out according to three parameters: 1) L / H -the ratio of the wing length to its width 2) Pt / r -the ratio of the pterostigma length to the length of the first segment of the radial vein 3) cell shape 2Rs , expressed by the ratio of the lengths of the segment of the radial sector and the sum of the lengths of the 4 and 5 segments of the basal vein (in A. viaticus and A. sabulosa), or the ratio of the lengths of the segment of the radial sector and the length of the 5 segment of the basal vein (in V. crabro), conventionally designated as 2Rs and 2Rs *. On the basis of the wing morphometric parameters of the right and left sides, the asymmetry of each individual was calculated. The ratio of the distances between the simple eyes and the faceted eye POL / OOL and the variability of body color (changes in the color and shape of the pattern) were also evaluated (Fig. 2).  Data on weather conditions are taken from the archive www.pogodaiklimat.ru/ (date of reference 03/13/2020). To calculate the hydrothermal coefficient (HTC), the following formula was used: where Σt͠ ≥10°C is the sum of average daily air temperatures higher than or equal to 10°C.
The sum of average daily temperatures was taken 5 years before the year in which the insects were caught. In order to indicate the value Σt͠ ≥10°С for 1893, Σt͠ ≥10°С were calculated for 1888, 1889, 1890, 1891, 1892, 1893 and Σt͠ ≥10°С was calculated from the obtained values. It is this average value of average temperatures for the previous 5 years that is designated Σt͠ ≥10°C in 1893. The values for 1956 and 2017 were calculated in the same way. АΣR (t͠ ≥10°С) is the amount of precipitation at an average daily air temperature above or equal to 10°C. The HTC value was calculated for the indicated years (1893,1956,2017). The graph of the secular variation of air temperature in Tomsk is taken from the literature (Sevast'yanov et al., 2000).
The Siberian silkworm Dendrolimus sibiricus Tschetverikov, 1908, according to literature data, was taken to review the changes in abundance in response to changing climatic conditions. To describe the transformation of the fauna of Western Siberia,

Results and Discussion
Analysis of meteorological data for 140 years allows us to make a conclusion about a systematic increase in the sum of average daily air temperatures during the warm period of the year (Fig. 3).  This graph shows a systematic increase in average annual temperatures with an increase in the amplitude of fluctuations between the maximum and minimum indicators in different years. Thus, it can be concluded that the tendency of climatic changes in this area to warming and aridization (the climate becomes hotter and drier). This climatic trend is typical not only for the Tomsk region. Our faunistic studies of recent years indicate changes in the composition of the West Siberian faunistic complex. Increasingly, on the territory of Western Siberia, we find typical representatives of the arid fauna (Central Asian and Ukrainian Journal of Ecology, 10(6), 2020 Mongolian). For example, in 2010 we discovered Podalonia alpina (Kochl, 1888) (Baghirov, 2010Chalybion turanicum (Gussakovskij, 1935 (Baghirov, 2011), in 2014-Evagetes trispinosus (Kohl, 1886 (Baghirov, 2014). All newly recorded species are southern and tend to arid and semi-desert biotopes. In other taxonomic groups, a similar picture can be observed. For example, in 2011 the Polygraphus proximus Blandford, 1894 was recorded for the first time (Baranchikov et al., 2011;Kerchev, 2014;Krivets et al., 2015a,b), later in 2014 in the Kemerovo region, and then in 2019 in the Tomsk region, a new species was discovered -Ips amitinus (Eichhoff, 1871) (Klimycheva, 2019). All this is the result of the expansion of the range due to changes in the climatic conditions of the Siberian region. Under the influence of global climate changes, the original range of all the listed species moved to the north of Russia, settling the western and northwestern regions from the Bryansk region to the Kola Peninsula. Now we find them in Siberia.
Another reaction to climate warming and drying can be observed in Dendrolimus sibiricus Tschetverikov, 1908. This species normally has a two-year life cycle (Rozhkov, 1965), but under conditions of a changed climate changes the normal physiological reaction and passes to a one-year life cycle. It should be noted that climatic factors only prepare the conditions for a breeding outbreak, but do not determine the dynamics of the process. If we turn to the history of the issue, then we find a clear correlation between outbreaks of numbers with dry and warm years. An outbreak of mass reproduction of the Siberian silkworm in Siberia was described in 1897 in Irkutsk. For the first time a Siberian silkworm was found in the Tomsk region in 1924 (Ioganzen, 1924;Burdavitcyn et al., 1957). Figure 5 shows that the average annual temperature during this period did not fall below 0 ° С. Then, in 1953Then, in -1955, described a grandiose wave of mass reproduction in the Tomsk region (Burdavitcyn et al., 1957). As we remember from Figures 3 and 4 in the period from 1893 to 1956 there is a sharp rise in temperature and a decrease in moisture content. Currently, biologists around the world are widely studying the effects of climate change. Entomologists have long been aware of the morphological reaction of the honey bee Apis mellifera Linnaeus, 1758 to an elevated temperature during ontogenesisthe wing venation changes until new veins and cells appear (these morphological structures for many insects are hallmarks of a species or even a genus). For 6 years, we studied the morphological changes of several species of insects within the same geographical point (collection materials for 120 years from 1893 to 2017) under the conditions of climate transformation. All studied species were analyzed by a complex of 7 characters (Figs. 2, 6). In this article, we present the results of three widespread wasp species from three different families: Pompilidae -Anoplius (Arachnophroctonus) viaticus (Linnaeus, 1758); Sphecidae -Ammophila sabulosa (Linnaeus, 1758); Vespidae -Vespa crabro Linnaeus, 1758. We have found that in all three species, a change in the symmetry of the head occurs -the distance between the posterior parietal ocelli increases and decreases between the parietal ocellus and the facet eye (Fig. 7). Also, in all three species, the proportion of the wing changes -the pterostigma (Pt) becomes shorter (Fig. 8).

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Insects under global warming. Review Ukrainian Journal of Ecology, 10(6), 2020 The venation of the wing of Anoplius (Ar.) viaticus (L.) acquired high variability (for example, the shape of the 2Rs cell). The shape of this cell in the modern population varies from trapezoidal to triangular on the leg. Along with other changes in Vespa crabro L., we noted a complication of the pattern on the abdomen and clypeus, as well as the replacement of light yellow in color with a dark yellow, almost orange color (Fig. 10).

Conclusion
Thus, for 120 years, we have been observing aridization of biotopes in Western Siberia and, as a result, we can observe the reaction of natural insect populations to climate changes in various manifestations -changes in the species composition, changes in the appearance of an insect, changes in the life cycle of a species.