CC BY
66
Biosystems
Diversity
ISSN 2519-8513 (Print) ISSN 2520-2529 (Online) Biosyst. Divers., 2019, 27(1), 76-84 doi: 10.15421/011912
Range of Pterostichus oblongopunctatus (Coleoptera, Carabidae) in conditions of global climate change
T. A. Avtaeva*, R. A. Sukhodolskaya**, A. V. Skripchinsky***, & V. V. Brygadyrenko****
*Complex Institute named after K. I. Ibragimov of the Russian Academy ofSciences, Grozny, Russia **Institute of Ecology of Tatarstan Academy of Sciences, Kazan, Russia ***North-Caucasus Federal University, Stavropol, Russia ****Oles HoncharDnipro National University, Dnipro, Ukraine
Article info
Received 03.01.2019
Received in revised form 10.02.2019
Accepted 12.02.2019
Complex Institute named after K I. Ibragimov of the Russian Academy of Sciences, M. Esambaev ave, 13, Grozny, 324024, Russia. Tel.: +88-712-222-676. E-mail: [email protected]
Research Institute for Problems of Ecology and Mineral Wealth Use ofTatarstan Academy of Sciences, Daurskaya st., 28, Kazan, 420087,Russia. Tel.: +795-031-526-19. E-mail: sukhodolskayaraisa@gmail. com
North-Caucasus Federal University, Pushkin st., 1, Stavropol, 355009, Russia. Tel.: +890-541-207-22. E-mail: [email protected]
Oles Honchar Dnipro National University, Gagarin av., 72, Dnipro, 49010, Ukraine. Tel.: +38-050-93-90-788. E-mail: [email protected]
Introduction
Avtaeva, T. A., Sukhodolskaya, R. A., Skripchinsky, A V., & Brygadyrenko, V. V. (2019). Range of Pterostichus oblongopunctatus (Coleoptera, Carabidae) in conditions of global climate change. Biosystems Diversity, 27(1), 76-84. doi:10.15421/011912
Using geodata technology, we conducted a bioclimatic modeling of the spatial distribution of the common palearctic ground beetle - Pterostichus oblongopunctatus (Fabricius, 1787). The range of comfort of the territories included in this species' range was obtained. We used the data on 510 sampling points, obtained as a result of the authors' field surveys and the data base of the GBIF global fund of biodiversity and 19 climatic parameters from the WorldClim open base and MaxEnt program. The results determined the factors which have the greatest impact on the current distribution of P. oblongopunctatus. The main climatic factors affecting the distribution of P. oblongopunctatus are average annual temperature, average 24-hour amplitude of temperature over each month, average temperature over the driest quarter, average temperature over the warmest quarter of the year, total of precipitations in the driest month of the year. We performed a prediction of possible change in the range by two scenarios (RCP 2.6 and RCP 8.5) for 2050 and 2070. Using QGIS program, we estimated the areas of the species' range, and compared them. According to the scenario RCP 2.6, by 2050, the range of the species will contract due to decrease in the territories with moderately continental climate, and by 2070, a restoration of the range would take place, for according to this scenario, the average annual temperature stabilizes. According to the scenario RCP 8.5, the range will contract by 2050 and will continue to decrease by 2070, for the concentration of CO2 continues to increase along with increase in average annual temperature. Climate changes can affect the life cycle of the beetle, its life expectancy and activity over the season. With changes in temperature, eggs and larvae of P. oblongopunctatus can be more vulnerable.
Keywords: global warming; climate change; range; ground beetles; modeling spatial distribution.
The problem of changes in ranges of common species in the conditions of global climate change is one most relevant problems of current ecology (Hijmans et al., 2005; Alonso-Carné et al., 2017). Climate change over the recent decades is an undeniable fact, which is proved by monitoring the rise in annual average temperature of air and ocean, melting of snow and ice on large areas and other climatic indicators. Climate change is a complex and multi-aspect phenomenon which affects practically all bioclimatic parameters (Zamolodchikov, 2016). Distribution of species is to a large extent determined by their ecological needs and peculiarities of spatial distribution of the most important factors limiting their global distribution (Musolin, 2007). They influence the spatial-temporal structure and system of interaction of organisms and their communities with the environment.
Modeling spatial distribution of individuals demonstrates the relationship between the presence of a particular species in the studied territory and the existing combination of bioclimatic factors. Modern geodata technologies and open data bases on bioclimatic parameters allow highly accurate prediction and mapping of the territories of distribution of biological objects, based on the knowledge of the limits of species' adaptability combined with spatial distribution of ecological factors of the environment (Panagos et al., 2017; Poggio et al., 2018). The following categories of reaction of insects to environmental changes are distingui-
shed: change in range, number, phenology, voltinism (number of generations per season), morphology, physiology and behaviour (Musolin, 2007).
The object of our study was Pterostichus oblongopunctatus (Fabricius, 1787), a transpalaearctic species distributed from Northern and Central Europe to the Caucasus and Siberia eastwards to Japan (Kry-zhanovskij et al., 1995; Hurka, 1996; Bousquet, 2003). It is one of commonest forest species of ground beetles in the south of the forest zone, in the forest-steppe zone and in the north of the steppe zone of Earasia (Kryshtak, 1956; Putchkov, 2018). In south of its range, its abundance reliably increases in conditions of 40-100% density of tree crowns, on loamy soils, at average levels of mineralization of soil. In the steppe zone of Ukraine the species is present in 19.5% of all studied forest ecosystems in the regions (Brygadyrenko, 2016a). According to Putchkov (2018), one of the factors which sometimes increase in the number of this forest species in the steppe zone is the humidity and shade regime (Brygadyrenko, 2015b, 2016b). Its occurrence in fields is related to intense irrigation, the species is common in old gardens and city parks of the forest and forest-steppe zones of Ukraine (Putchkov, 2018).
According to Putchkov (2018), in Ukraine, females of the species with eggs have been "recorded in April-May. Larvae develop from May to June. Appearance of young beetles was observed in August-September. Imagoes overwinter. Zoophage, consumes lepidopterans related to soil (Noctuidae, Geometridae, Cydia), Staphylinidae (genus Philon-thus), larvae and pupae of scarabs (Scarabaeidae), darkling beetles
(Tenebiionidae), click beetles (Elateiidae), worms and their cocoons, small slugs. They have been observed to eat trama of mushrooms and larvae of Mycetophilidae, which live in them, and also sporadically -fruits of strawberry". This species is distinct in the high diversity of its diet in undamaged forest ecosystems both at the larva and imago stages of development (Schelvis & Siepel, 1988; Komarov & Brygadyrenko, 2011). Duration of reproductive period is affected by the latitude of the inhabited location, absolute height above sea level, anomalous weather conditions and habitats (Van Schaick Zillesen, 1985; Sharova & Filippov, 2003; Matalin, 2006). In the forest-steppe, P. oblongopunctatus breeds from early April to late July (Sharova & Denisova, 1997); in broad-leaved forests, the reproductive period lasts from late April to late July (Heerdt et al., 1976); in coniferous-broad leaved forests - from early May to Mid July; in the northern taiga - from early June to early July (Sharova & Filippov, 2003). In highland areas of the Chechen Republic P. oblongopunctatus is active from early May to late September.
This species is quite a convenient practical object for physiological studies: there are studies of the effect of nickel on its own and in combination with chlorpyrifos and heightened temperature on larvae (Bednar-ska & Laskowski, 2009) and imagines of this species (Bednarska & Laskowski, 2007; Bednarska et al., 2009; Bednarska & Kaszowska, 2014). Toxicological studies on this species led to a number of important conclusions being drawn on the toxicology of the pollutants, applicable for other species of ground beetles and insects in general (Lagisz et al., 2002; Bednarska et al., 2010a, 2010b, 2012, 2016, 2017; Bednarska & Stachowicz, 2012; Simon et al., 2016). Work on different aspects of po-pulational diversity on genetic (Siepel, 1988; Den Boer et al., 1992; Lagisz et al., 2010) and morphological (Emetz, 1986; Schwerk & Jaskula, 2018) levels allow one to state that P. oblongopunctatus is one of the best studied species of ground beetles.
P. oblongopunctatus is a typical element of urban forest plantations and anthropogenically transformed territories (Grechanichenko & Guseva, 2000; Magma et al., 2008; Gardiner & Harwood, 2017). Thus, the studied species of ground beetle is a practical object for modeling changes in the range in conditions of global climate changes, one of the priority factors which affects natural populations in XXI century (Kotze et al., 2011). It was experimentally determined that temperature is an important limiting factor for this species of ground beetle (Thiele, 1975, 1977; Bednarska & Laskowski, 2007).
The objective of this study was to assess the distribution of P. oblongopunctatus in the context of changes in global climate conditions by using ecological-climatic modeling.
Material and methods
Material for this study was collected in the field by the authors (120 sites), and also the data obtained from the GBIF open data base (390 sites). Collecting in the field was performed in different years on the territory of Russia (Tatarstan, Chechen Republic, Sverdlov Oblast, west slope of the Urals, Perm Krai) and Ukraine (Table 1).
Table 1
Places where P. oblongopunctatus was collected for the analysis of changes in the range
Country, region_Oblast, region_
Ukraine Kyiv, Cherkasy, Poltava, Kharkiv, Dnipropetrovsk
Zaporizhia, Mykolaiv, Donetsk Oblasts Tatarstan Kazan, Rybno-Slobodsky district, Apastovsky district,
Volga-Kama National Reserve, Laishevsky district, Zelenodolsky district, Vysokogorsky district, Verhneuslonsky district, Island biotopes on the Volga Sverdlovsk Oblast Ekaterinburg, Pripyshminskiye Bory National Park,
Zyrianovsky district, Visim National Natural Biosphere Reserve Kemerovo Oblast Kemerovo Mari El Republic Mariy Chodra National Park
Chechen Republic Vedensky and Sharoysky districts_
The beetles were sampled using pitfall traps and soil samples, and also collected manually, mostly in mesophilous habitats in flood-plain, broad-leaved and coniferous forests; more rarely - in shrub and mea-
dow communities. This allowed broad data to be obtained which reflected different aspects of ecology and biology of this species. In total over 11 thousand imagoes of P. oblongopunctatus were collected.
For bioclimatic modeling (Molyneux et al., 2013), WorldClim data of global base of climatic data (www.worldclim.org) were used: 19 bioclimatic variables with spatial resolution of 30 seconds. For the analysis of changes in the range of P. oblongopunctatus, we used bioclimatic data for the years 1960-1990 (version 1.4 WorldClim), bioclimatic data for 1970-2000 (version 2 WorldClim), and also two scenarios of 'low emissions" (RCP 2.6) and "high emissions" (RCP 8.5) out of four predicted scenarios, RCP 2.6, RCP 4.5, RCP 6.0, RCP 8.5, which became known as "representative concentrations pathways" (www.global-climate-change.ru). RCP correlate with a broad spectrum of probable changes in anthropogenic emissions of greenhouse gases and are used for demonstrating their concentration in the atmosphere. The scenario of 'low emissions", RCP 2.6, includes constant and ambitious actions orientated towards reduction of anthropogenic emissions of greenhouse gases. Trajectories of RCP 2.6 have a large variation of annual changes in temperature, which leads to slowing and even stopping its growth after 2050. The reasons for such slowing are determined by the dynamic of CO2 concentration, which in turn is related to significant absolute decrease in anthropogenic emissions of greenhouse gases, starting from 2020 in RCP 2.6.
RCP 8.5 characterizes a situation when global economic development remains associated with use of fossil fuel without any measures of climatic policy being taken. During the realization of RCP 8.5, by 2100, the concentration of CO2 will reach 936 ppm, and the average global temperature will reach 16.8 °C. In other words, warming over 2010-2100 will equal 2.4 °C. During RCP 2.6, by 2100, concentration of CO2 will equal 421 ppm, and growth of global temperature - only 0.1 °C. According to the prognosis of the Intergovernmental Panel on Climate Change (IPCC), on the trajectory of RCP 8.5, annual increase in temperature will vary within 0.03-0.05 °C.
The mean value of global temperature in 1850-1990 (pre-industrial level) equaled 13.6 °C (Fick & Hijmans, 2017). Therefore, after achieving the goals of the Paris Agreement, mean global temperature should not exceed 15.6 °C. According to the prognosis on the CMIP 5 ensemble of models, global temperature will reach 15.6 °C in2042 according to RCP 8.5; in 2056 according to RCP 4.5; and in 2062 according to RCP 6.0. Only scenario RCP 2.6 appears to correspond to the goal of the Paris Agreement.
Zamolodchikov (2016) suggested a statistical model of the dynamic of average annual global temperature which combines logarithmic effect of carbon dioxide concentration growth and contribution of the climatic cycles. Parameters of the model are determined according to known data of instrumental measurements over 1850-2010. The model proves the reliable presence of two cyclic process of periodicity in the dynamics, equaling 10.5 and 68.8 years. According to the model, maxima of this cycle should be observed in the years 1944, 2012, 2081. The amount of annual change in temperature was positive and was within 0.08-0.09 °C in 1989-2000. This particular period was characteristic of the fastest factual growth of global temperature. However, subsequently, the annual change due to the 68.8-year cycle started to decrease, and in 2005, it fell to 0.05 °C, and from 2012 became negative, i.e. decreasing the tempi of global temperature growth. According to this model, exceeding the threshold of 2 °C will occur in 2061 at RCP 8.5 and in 2075 at RCP 6.0.
We used QGIS 3.4.3 (Quantum GIS, 2019) for working with layers, calculation of points number in the polygons, assessment of area of the predicted ranges. Predicting potential range was made using Maxent 3.4.1 program (Phillips, S. J., Dudik, M., & Schapire, R. E., 2017: MaxEnt software for modeling species niches and distributions (Version 3.4.1), http://biodiversityinformatics.amnh.org/open_source/maxent).
Results
Modeling in medium MaxEnt demonstrated that the greatest impact on the spatial distribution of P. oblongopunctatus was caused by the following bioclimatic variables of Bioclim (percentage of contribution of each variable is provided in Table 2):
- BIO 01 - average annual temperature;
- BIO 02 - average 24-hour amplitude of temperature for each month;
- BIO 09 - average temperature of the driest quarter;
- BIO 10 - average temperature of the warmest quarter of the year;
- BIO 14 - total of precipitations in the driest month of the year. For the assessment of the obtained model, a testing selection was
created, which included 25% of the species presence points. The results are presented in logistical format which provides 0 to 1 assessment of the probability of this species inhabiting a particular geographic point. The map of potential distribution of the studied species demonstrates how different territories are appropriate for it according to their climatic characteristics.
An important indicator of the model's reliability is Area Under the Curve (AUC). AUC is assessment of the ability of the model to demonstrate presence of the species in a geographic point, where it could live with a high percentage of probability. In our case, AUC equals 0.975. That means a 97.5% probability that the species will actually live where is predicted to live. The model of distribution of P. oblongopunctatus
corresponds to the current knowledge of the places where the species lives (Fig. 1).
Table 2
Percentage of contribution and importance
with permutation of the studied bioclimatic parameters
Bioclimatic parameters Contribution of bioclimatic parameter, % Importance of the parameter during permutation, %
BIO 14 52.6 1.7
BIO 01 14.3 11.4
BIO 11 10.3 0.0
BIO 09 7.4 1.9
BIO 02 6.0 18.1
BIO 04 3.2 1.7
BIO 13 2.6 8.8
BIO 10 1.2 43.6
BIO 19 0.4 5.5
Fig. 1. Map of potential distribution of P. oblongopunctatus: colour scale of the comfort range: dark blue - unfavourable (0-0.03); light blue -favourable (0.04-0.12); yellow and green - favourable territories (0.13-0.60); red and orange - most favourable territories (0.61-0.98)
less
Using QGIS, we performed an analysis of the number of points in the polygons for each factor. The number of points in each polygon was obtained from the table and on their basis diagrams were built. Out of 510 points, 390 are in the range of average annual temperature 4-10 °C (Fig. 2). Perhaps, this is most favourable temperature for this species. During the analysis of average 24-hour amplitude of temperature, 384 points were obtained in range of 7-10 °C (Fig. 3).
Fig. 2. Distribution of records points in P. oblongopunctatus (abscissa axis) depending on average annual air temperature (ordinate axis, °C)
Analysis of average temperature of the warmest quarter demonstrates that 437 points are in the range of 9.0-10.5 (Fig. 4). At the analysis of total of precipitations of the driest month of the year, 347 points are in the range of 25-50 mm (Fig. 5).
Fig. 3. Distribution of record points in of P. oblongopunctatus (abscissa axis) depending on the average 24-hour amplitude of temperature (ordinate axis, °С) for each month
14.S-14.9 14.0-14.5 13.5-14.0 13.0-13.5 12.5-13.0 12.0-12.5 11.5-12.0 11.0-11.5 10.5-11.0 10.0-10.5 9.5-10.0 9.0-9.5 8.5-9.0 8.0-8.5 7.5-8.0 7.0-7.5
Fig. 4. Distribution of records points in P. oblongopunctatus (abscissa axis) depending on the average temperature of the warmest quarter of the year (ordinate axis, t)
Fig. 5. Distribution of record points in P. oblongopunctatus (abscissa axis) depending on the total amount of precipitations (mm) over the driest month of the year
In QGIS program, using a calculator of fields, we estimated the area of current range, area of ranges predicted for 2050 and 2070 according to the scenarios RCP 2.6. and RCP 8.5 (Fig. 6).
Fig. 6. Change in the area of range (km ) of P. oblongopunctatus : striped column - scenario RCP 2.6, grey column - scenario RCP 8.5
According to the first scenario, beginning with the second half of the XX century to 2020, the content of CO2 will increase, as well as the global average annual temperature. After 2020, anthropogenic emissions will decrease, and the average annual temperature will stabilize. According to the model we obtained, the area of range of P. oblongopunctatus will decrease until 2050 (Fig. 7), and by 2070, due to decrease in CO2 concentration and reduction of average annual increase in temperature, the range will begin to recover (Fig. 8).
Fig. 7. Change in the area of range by 2050 according
to the scenario RCP 2.6: lilac tint - current range, diagonally hatched ( yellow ) areas - range in year 2050
Fig. 8. Change in the range by 2070 according to scenario RCP 2.6: plain tint - range as at 2070, hatched areas - range in 2050
According to scenario RCP 8.5 (Fig. 9), the range of the species looks different - the concentration of CO2 and mean annual temperature will continue to increase, which will lead to reduction of the range and shift of the borders of the comfort zone to areas with a mild maritime climate. Areas with continental climate become unfavourable for survival of P. oblongopunctatus, which is perhaps related to the sharpening of temperature differences and reduction of the amount of precipitations.
Fig. 9. Change in area of range by 2070 according to RCP 8.5: Plain tint - range as at 2050, hatched areas - range as at 2070
According to scenario RCP 2.6, by 2050 in North America, the range will significantly contract inside the continent, but increase in the islands (Fig. 10). The same picture can be seen in the islands of the coast of the Pacific Ocean in Asia (Fig. 11). According to scenario RCP 8.5, P. oblongopunctatus will not live in these islands.
We used the MaxEnt program for assessment of the contribution of mean monthly maximum and minimum temperatures to the realization of life cycle of P. oblongopunctatus, particularly jackknife function. The result ofthe analysis is demonstrated in bar charts (Fig. 12, 13).
Fig. 10. Range in North America at present and in 2050 according to scenario RCP 2.6
Fig. 11. Range on islands of Pacific Ocean coast: prediction for 2050 according to scenario RCP 2.6
Fig. 12. Bar chart of contribution of mean monthly maximum temperature on life cycle of P. oblongopunctatus, parameter of factor significance (abscissa axis); mean maximum temperature from January to December (ordinate axis)
Blue bars indicate the role of the variable in development of model. The shorter is the blue bar, the larger amount of unique information a variable contains. In this case, the most significant are maximum temperatures of the cold period: January-March, September-December. A similar picture is observed for minimum temperatures.
This is supported by the fact that the development of the gonads is significantly affected by daylight hours and temperature, due to which a strong correspondence to particular seasonal periods is observed. The go-nads of male P. oblongopunctatus become mature in the conditions of short days, i.e. in autumn before wintering and during wintering, and gonads of females - after short days become long, i.e. in spring right after wintering (Thiele, 1966; Thiele & Konen, 1975; Kreckwitz, 1980).
Fig. 13. Bar chart of contribution of every month's mean minimum
temperature to life cycle of P. oblongopunctatus, parameter of factor significance (abscissa axis); mean maximum temperature from January to December (ordinate axis)
We described the seasonal dynamic and life cycle of P. oblongopunctatus which lives in the conditions of mountains of Chechnia. Seasonal activity was observed from early May. Females with unlay ed eggs and generative males can be found from the first decade of May to late July, becoming most numerous in the second decade of June. In the third decade of May, postgenerative individuals were found in the selections in small numbers. The first juvenile specimens began to sampled in the traps in the third decade of June. The peak of juvenile individuals occurred in the second decade of July. Along with generative, postgenerative and juvenile individuals, in late July, the traps sampled immature individuals, the number of which remains stable until late September. In the autumn selections, juvenile and postgenerative individuals there were also observed. Thus, the life cycle of P. oblongopunctatus in the conditions of mountains of Chechnia is realized as a one-year cycle with spring-summer breeding.
Discussion
Current distribution. Mapping the ranges of this ground beetles was performed only for certain countries: the Netherlands and Great Britain (Turin et al., 1977; Luff et al., 1989). The ecology of P. oblongopunctatus in Fennoscandia and Denmark, according to Lindroth (1986), corresponds to its ecology in Central Europe. This famous author emphasizes the eurotopicity of P. oblongopunctatus, as a forest species: "euryto-pic woodland species, occurring in both deciduous and coniferous forests, usually in light stands on moderately dry, mainly sour humus soil". Lindroth (1986) also accents that in the conditions of sufficiently humid and cool climate, the species comes out of the forest canopy to open territories: "In the Atlantic climate of West Norway also in open habitat". According to Koivula et al. (1999), the species statistically reliably reduces its abundance in response to decrease in the layer of forest litter in Central Finland. P. oblongopunctatus was found to be a common species in natural spring deposits in Late Holocene in South-Central Sweden (Hellqvist, 2014).
In conditions of sufficient moisture in Great Britain, P. oblongo-punctatus lives in all kinds of soil, often under bark. According to Lindroth (1974): it is found in England, in the north up to Yorkshire (lacking in the S.E.), Wales, Scotland, Ireland. "Inhabits woodland and forests, occurring over much of Britain. It has also been recorded from Ireland. Its distribution is rather patchy, with four or five apparent centres of distribution, and it is very local in Ireland. It generally shows a preference for rather open woods on dry soils", according to Luff (1992).
According to Freude et al. (2004), in the forests of Central Europe, the species is frequently found. P. oblongopunctatus is mentioned for the entire territory of Ukraine (Putchkov, 2011, 2012). In Southern Europe, the range of species is studied quite well (Arndt et al., 2011; Brigic et al., 2014). Gueorguiev (2007) provided mention for only 3 places in Albania, Hieke & Wrase (1988) - for 25 in Bulgaria. Later, Gueor-
guiev & Gueorguiev (1995) indicate records of this species in 13 of 22 regions. Further to south, this species occurs sporadically: Hristovski & Gueorguiev (2015) for the first time recorded this species in 3 from 93 geographical regions of Macedonia, Serrano (2013) recorded P. oblongopunctatus only in 1 (Pyrenees) out of 23 geographical regions of Spain.
In the central part of its range, in the Czech and Slovak Republics, P. oblongopunctatus is "very common in all types of forests; lowlands to mountains, frequent in hills" (Hurka, 1996). This species is abundant in the forests of South Moravia (Sejnohova, 2006). The species dominates in the forests of Poland (Sklodowski, 2006, 2014; Ulrich & Zalew-ski, 2006; Zalewski & Ulrich, 2006; Kwiatkowski, 2011; Sklodowski & Garbalinska, 2011; Zalewski et al., 2012). There numerous reports of finding this species in Lithuania (Tamutis et al., 2011). According to contemporary data (Putchkov, 2018), the species in the Ukrainian part of the range is common "in Polesie and Forest-Steppe, more often is found in forests of different types, near water bodies, in humid areas, in woodland belts, oak forests, groves, sometimes in meadows near the woodlands. In mountains, it reaches the subalpine zone. In the Steppe, it lives only in bairak forests and floodplain tree-shrub biotopes". In the Steppe zone of Ukraine, this is one of the commonest species in differ-rent types of forests, and beyond the borders of forest communities, P. oblongopunctatus is recorded rarely along river banks, meadows near woodland edges, in thinned out city parks, though it prefers parts of forests with typical mesophilous microclimatic conditions and developed forest litter (Brygadyrenko, 2016). The species is common in forest ecosystems of Moldova (Nekuliseanu & Matalin, 2000).
Kryzhanovskij (1983) indicates this species as being commonest in both the taiga and mixed forest zones. Kryzhanovskij et al. (1995) indicate the following distribution for this species in Russia: the northern Russian plain, the central part of the Russian plain, the southern Russian plain, the Caucasus Major, the Urals, northern West Siberia, the middle stretch of West Siberia, southern West Sibera, The Altais-Sayan Mt. Land, Middle Siberia, and Transbaikalia. In the zone of broad-leaved forests it is one of the most numerous species of ground beetles (Sharova, 1981; Sergeeva & Grunthal, 1988; Soboleva-Dokuchaeva, 1995; Sharova & Denisova, 1997; Zherebcov, 2000; Belskaya & Zolotarev, 2017). In the Volga region, it is found quite often, but is rare in the south (Kaljuzhnaja et al., 2000). In Ciscaucasia, it lives in forests of mesophilous type (Kryzhanovskij, 1983; Sigida, 1993); in Republic of Adygheya it is common in forests, and found more frequently in humid places (Zamotajlov & Nikitsky, 2010). Babenko & Eremeyeva (2007) report that this species as dominant in upland meadows outcide the city of Kemerovo. In the Far East, a whole group of closely related species of groundbeetles is distributed (Kryzhanovskij et al., 1995).
Probable change in the range. P. oblongopunctatus is a polyzonal species (Putchkov, 2018), therefore the shift of its range towards the north in the XXI century is perhaps related to the change in density of distribution of forest ecosystems. In QGIS program, we obtained maps which depict the range of comfort for P. oblongopunctatus at the current moment (Fig. 14), prognosis according to the scenario RCP 2.6 for year 2050 (Fig. 15) and prognosis for 2070 (Fig. 16). At the same time, using GIS-technologies allows recording absence or shifting of the range.
If one analyzes which of the stages of the seasonal dynamics (stages of ontogenesis of P. oblongopunctatus) can become the most vulnerable, in the south the most probable will be death of eggs and larvae during spring or summer droughts. In the territory of Ukraine, the species was reported throughout the vegetative season: from early spring to late autumn (Putchkov, 2018). Spring breeding and summer development of larvae which have thinner coats and which are less able to move quickly, increases the probability that this species will suffer a negative effect from the perspective of probable summer droughts in the south part of its range. Perhaps, death of larvae in the litter can be caused by forest fires.
In the north part of the range, probably the border of distribution range of P. oblongopunctatus will be the northernmost parts of forest ecosystems discovered to act as biological communities. One can presume that changes in mean annual temperature in the north part of the range can have an effect on the period of activity of the studied species. Late
spring and summer breeding can be replaced by spring and even early spring breeding, and the beetle's activity period can be prolonged. The duration of the development of each stage of ontogenesis in Carabidae is influenced by a number of factors: temperature regime of soil and terrestrial layer of air, moisture, daylight hours, amount and quality of food, its accessibility, etc. (Thiele, 1975, 1977; Korolev & Brygadyrenko, 2014; Brygadyrenko, 2015a; Brygadyrenko & Korolev 2015).
Fig. 14. Current range of comfort for P. oblongopunctatus: red and orange - most favourable territories, yellow and green - favourable territories, light blue - less favourable, dark blue - unfavourable territories
Fig. 15. Range of comfort for P. oblongopunctatus in 2050 according to scenario RCP 2.6: for notes see Fig. 12
Fig. 16. Range of comfort for P. oblongopunctatus in 2070 according to scenario RCP 2.6: for notes see Fig. 12
Usually, the fastest development occurs among eggs, larvae of first age and pupae, whereas the development of larvae of older ages requires much more time. Even in the most unfavourable conditions, a minimum
temporal threshold of pre-imago development exists, which among the majority of ground beetles of the temperate zone equals 1.0-1.5 months. One should also take into account the fact that the development of gonads of many species is controlled by photoperiod and therefore is closely associated with certain seasonal periods. Shift in range can lead to disorders of trophic relations, decrease in accessibility of the food base.
Analysis of models in range ofP. oblongopunctatus revealed that the largest area of the range corresponds to the period 1960-1990. The zone of highest comfort is in the range of 0.85-0.92 with a maximum of 1 in the south-east coast of the central and southern part of Sweden, in south-west central and south part of Finland, in north-west countries of the Baltic, in the north of Poland and Germany, in the territory of Denmark, north-west coast of the Black Sea, eastern coasts of England and south of Kamchatka. The zone of heightened comfort for P. oblongopunctatus is mainly in the territory with mild maritime climate.
The model developed using contemporary bioclimatic data differs in the smaller area of the range, maximum values of the range of comfort at the territory of occurrence shift by 0.77-0.69. According to obtained contours, the predicted ranges in 2050 and 2070 according to scenario RCP 8.5 are significantly different from the predicted ranges according to scenario RCP 2.6. Large territories of Poland, Hungary, Romania, Ukraine become excluded from the zone of comfort of P. oblongopunctatus. According to scenario RCP 8.5, area of zone of comfort decreases on islands, whereas according to scenario RCP 2.6, an opposite effect is observed. According to scenario CRP 2.6, by 2050, a shift of the range will be observed towards the west, and also the area will decrease by 2070, when CO2 emissions will practically stop, mean annual temperatures will have increased by 0.1 °C, and the range of the studied species will begin to recover (Fig. 14).
Conclusions
Because the scenarios do not take into account biotic factors, food base and anthropogenic effect, the confident prognosis of change in the range is impossible. Also, it is impossible to consider all populations at the current stage of existence of the species and predict under which of the climate change scenarios they will develop in the future. However, the data we received indicate that the temperature regime is the main factor which directly or indirectly affects the distribution of this ground beetle. We hope that similar, though far from numerous, studies on the impact of global climatic changes on the distribution of certain species of ground beetles will help to conserve these zoophages which are important for natural ecosystems.
We express our gratitude to N. I. Eeremeeva, Grand PhD in biology (Kemerovo State University), S. L. Esiunin, Grand PhD in biology (Perm State University), V. I. Matveev, assistant lecturer (Mari State University) for provision of materials on distribution ofP. oblongopunctatus.
References
Alonso-Carné, J., Estrada-Peña, A., & García-Martín, A. (2017). Análisis comparativo de la información de temperatura de las bases de datos WorldClim, ECA&D y MODIS en el Paleártico Occidental. Geographicalia, 69, 1. Arndt, E., Schnitter, P., Sfenthourakis, S., & Wrase, D. (Eds.). (2011). Ground
beetles (Carabidae) of Greece. Pensoft, Sofia - Moscow. Babenko, A. S., & Eremeyeva, N. I. (2007). Osobennosti naseleniya zhuzhelits urbanizirovannykh territoriy v usloviyakh sibirskikh gorodov [Peculiarities of the communities of ground beetles of urbanized areas in the conditions of Siberian cities]. Vestnik Tomskogo Gosudarstvennogo Universiteta, Biolo-giya, 1, 5-17 (in Russian). Bednarska, A. J., & Kaszowska, J. (2014). Combined effect of nickel and chlorpy-rifos on the ground beetle Pterostichus oblongopunctatus. Ecotoxicology and Environmental Safety, 108, 242-248. Bednarska, A. J., & Laskowski, R. (2007). Effects of nickel and temperature on the ground beetle Pterostichus oblongopunctatus (Coleoptera: Carabidae). Ecotoxicology, 17(3), 189-198. Bednarska, A. J., & Laskowski, R. (2009). Environmental conditions enhance toxicant effects in larvae of the ground beetle Pterostichus oblongopunctatus (Coleoptera: Carabidae). Environmental Pollution, 157(5), 1597-1602.
Bednarska, A. J., & Stachowicz, I. (2012). Costs of living in metal polluted areas: Respiration rate of the ground beetle Pterostichus oblongopunctatus from two gradients of metal pollution. Ecotoxicology, 22(1), 118-124.
Bednarska, A. J., Brzeska, A., & Laskowski, R. (2010a). Two-phase uptake of nickel in the ground beetle Pterostichus oblongopunctatus (Coleoptera: Cara-bidae): Implications for invertebrate metal kinetics. Archives of Environmental Contamination and Toxicology, 60(4), 722-733.
Bednarska, A. J., Gerhardt, A., & Laskowski, R. (2010b). Locomotor activity and respiration rate of the ground beetle, Pterostichus oblongopunctatus (Coleoptera: Carabidae), exposed to elevated nickel concentration at different temperatures: Novel application of multispecies freshwater biomonitor®. Ecotoxicology, 19(5), 864-871.
Bednarska, A. J., Laskowski, R, Pyza, E., Semik, D., Swi^tek, Z., & Woznicka, O. (2016). Metal toxicokinetics and metal-driven damage to the gut of the ground beetle Pterostichus oblongopunctatus. Environmental Science and Pollution Research, 23(21), 22047-22058.
Bednarska, A. J., Portka, I., Kramarz, P. E., & Laskowski, R. (2009). Combined effect of environmental pollutants (nickel, chlorpyrifos) and temperature on the ground beetle, Pterostichus oblongopunctatus (Coleoptera: Carabidae). Environmental Toxicology and Chemistry, 28(4), 864-872.
Bednarska, A. J., Stachowicz, I., & Kurianska, L. (2012). Energy reserves and accumulation of metals in the ground beetle Pterostichus oblongopunctatus from two metal-polluted gradients. Environmental Science and Pollution Research, 20(1), 390-398.
Bednarska, A. J., Swi^tek, Z. M., Paciorek, K., & Kubinska, N. (2017). Effect of cadmium bioavailability in food on its compartmentalisation in carabids. Ecotoxicology, 26(9), 1259-1270.
Belskaya, E. A., & Zolotarev, M. P. (2017). Changes in the size structure of carabid communities in forest ecosystems under technogenic transformation. Russian Journal of Ecology, 48(2), 152-160.
Bousquet, Y. (2003). Tribe Pterostichini. In: Löbl, J., & Smetana, A. (Eds.). Catalogue of Palearctic Coleoptera. Vol. 1. Archostemata - Myxophaga - Ade-phaga Apollo Books, Stenstrup. Pp. 462-521.
Brigic, A., Starcevic, M., Hrasovec, B., & Elek, Z. (2014). Old forest edges may promote the distribution of forest species in carabid assemblages (Coleoptera: Carabidae) in Croatian forests. European Journal of Entomology, 111(5), 715-725.
Brygadyrenko, V. V. (2015a). Evaluation of the ecological niche of some abundant species of the subfamily Platyninae (Coleoptera, Carabidae) against the background of eight ecological factors. Folia Oecologica, 42(2), 75-88.
Brygadyrenko, V. V. (2015b). Influence of moisture conditions and mineralization of soil solution on structure of litter macrofauna of the deciduous forests of Ukraine steppe zone. Visnyk of Dnipropetrovsk University. Biology, Ecology, 23(1), 50-65.
Brygadyrenko, V. V. (2016a). Evaluation of ecological niches of abundant species of Poecilus and Pterostichus (Coleoptera: Carabidae) in forests of steppe zone of Ukraine. Entomologica Fennica, 27(2), 81-100.
Brygadyrenko, V. V. (2016b). Influence of litter thickness on the structure of litter macrofauna of deciduous forests of Ukraine's steppe zone. Visnyk of Dnipropetrovsk University. Biology, Ecology, 24(1), 240-248.
Brygadyrenko, V. V., & Korolev, О. V. (2015). Morphological polymorphism in an urban population of Pterostichus melanarius (Illiger, 1798) (Coleoptera, Carabidae). Graellsia, 71(1), e025.
Den Boer, P. J., Vermeulen, R., & Szyszko, J. (1992). Spreading the risk of extinction by genetic diversity in populations of the carabid beetle Pterostichus oblongopunctatus F. (Coleoptera, Carabidae). Netherlands Journal of Zoology, 43(3-4), 242-259.
Emetz, V. M. (1986). Changes in spatial-distribution of adults in a population of the ground beetle Pterostichus oblongopunctatus F (Coleoptera, Carabidae) under the influence of recreation. Zhurnal Obshchei Biologii, 47(1), 125-127.
Fick, S. E., & Hijmans, R J. (2017). WorldClim 2: New 1-km spatial resolution climate surfaces for global land areas. International Journal of Climatology, 37(12), 4302-4315.
Freude, H., Harde, K.-W., Lohse, G. A. (2004). Die Käfer Mitteleuropas. Band 2. Adephaga. 1. Carabidae (Laufkäfer). Elsevier, Spektrum Akademischer Verlag, Heidelberg, Germany.
Gardiner, M. M., & Harwood, J. D. (2017). Influence of heavy metal contamination on urban natural enemies and biological control. Current Opinion in Insect Science, 20, 45-53.
Grechanichenko, T. E., & Guseva, N. A. (2000). Specific features of carabid populations (Coleoptera, Carabidae) in forest biotopes of central forest-steppe. Zoologichesky Zhurnal, 79(5), 548-555.
Guéorguiev, B. (2007). Annotated catalogue of the carabid beetles of Albania (Coleoptera: Carabidae). Pensoft Publishers, Sofia-Moscow, Bulgaria.
Guéorguiev, V. B., & Guéorguiev, B. V. (1995). Catalogue of the ground-beetles of Bulgaria (Coleoptera: Carabidae). Pensoft Publishers, Sofia-Moscow, Bulgaria.
Heerdt, P. F., Blokhuis, B., & Haafen, C. (1976). The reproductive cycle and composition of a population of Pterostichus oblongopunctatus (Fabricius) in the Netherlands (Coleoptera: Carabidae). Tjdschrft Voor Entomologie, 119(1), 10-13.
Hellqvist, M. (2014). Analysis of coleopterans from Late Holocene natural spring deposits in South-Central Sweden. Quaternary International, 341, 184—194.
Hieke, F., & Wrase, D. W. (1988). Faunistik der Laufkafer Bulgariens (Coleoptera, Carabidae). Deutsche Entomologische Zeitschrift, 35(1-3), 1—171.
Hijmans, R. J., Cameron, S. E., Parra, J. L., Jones, P. G., & Jarvis, A. (2005). Very high resolution interpolated climate surfaces for global land areas. International Journal of Climatology, 25, 1965—1978.
Hristovski, S., & Gueorguiev, B. (2015). Annotated catalogue of the carabid beetles of the Republic of Macedonia (Coleoptera: Carabidae). Zootaxa, 4002(1), 1—190.
Hurka, K. (1996). Carabidae of the Czech and Slovak Republics. Print Centrum, a. s., Zlin, Czech Republic.
Kaljuzhnaja, N. S., Komarov, E. V., & Cherezova, L. B. (2000). Zhestkokrylye nasekomye (Insecta, Coleoptera) Nizhnego Povolzhja [Coleopterous insects (Insecta, Coleoptera) of Lower Volga area]. Volgograd, Russia (in Russian).
Koivula, M., Punttila, P., Haila, Y., & Niemela, J. (1999). Leaf litter and the small-scale distribution of carabid beetles (Coleoptera, Carabidae) in the boreal forest. Ecography, 22(4), 424—435.
Komarov, O. S., & Brygadyrenko, V. V. (2011). Trophic preferences of Pterostichus oblongopunctatus (Coleoptera, Carabidae) in conditions of south forest-steppe. Visnyk of Dnipropetrovsk University, Biology, Ecology, 19(1), 41—50.
Korolev, О. V., & Brygadyrenko, V. V. (2014). Influence of individual variation in the trophic spectra of Pterostichus melanarius (Coleoptera, Carabidae) on the adaptation possibilities ofits population. Folia Oecologica, 41(1), 34—43.
Kotze, D. J., Brandmayr, P., Casale, A., Dauffy-Richard, E., Dekoninck, W., Koivula, M. J., Lovei, G. L., Mossakowski, D., Noordijk, J., Paarmann, W., Pizzolotto, R., Saska, P., Schwerk, A., Serrano, J., Szyszko, J., Taboada, A., Turin, H., Venn, S., Vermeulen, R., & Zetto, T. (2011). Forty years of carabid beetle research in Europe — from taxonomy, biology, ecology and population studies to bioindication, habitat assessment and conservation. ZooKeys, 100, 55—148.
Kryshtal, O. P. (1956). Entomofauna gruntu ta pidstylky v dolyni serednioji techiji riky Dnipro [Entomofauna of soil and litter in the valley of Dnipro river midstream]. Kyiv State University, Kyiv (in Ukrainian).
Kryzhanovskij, O. L. (1983). Zhuki podotrjada Adephaga: Semejstva Rhysodi-dae, Trachypachidae; semejstvo Carabidae (Vvodnaja chast' i obzor fauny SSSR) [The beetles of the suborder Adephaga: Families Rhysodidae, Trachypachidae, Carabidae (Introduction and a review of the USSR fauna)]. Nauka, Leningrad (in Russian).
Kryzhanovskij, O. L., Belousov, I. A., Kabak, I. I., Kataev, B. M., Makarov, K. V., & Shilenkov, V. G. (1995). A checklist of the ground-beetles of Russia and adjacent lands (Insecta, Coleoptera, Carabidae). Pensoft Publishers, Sofia, Bulgaria.
Kwiatkowski, A. (2011). Assemblages of carabid beetles (Coleoptera, Carabidae) in humid forest habitats of different stages of succession in the Puszcza Knyszyn-ska Forest (Northeastern Poland). ZooKeys, 100, 447—459.
Lagjsz, M., Kramarz, P., Laskowski, R., & Tobor, M. (2002). Population parameters of the beetle Pterostichus oblongopunctatus F. from metal contaminated and reference areas. Bulletin of Environmental Contamination and Toxicology, 69(2), 243—249.
Lagisz, M., Wolff, K., Sanderson, R A., & Laskowski, R. (2010). Genetic population structure of the ground beetle, Pterostichus oblongopunctatus, inhabiting a fragmented and polluted landscape: Evidence for sex-biased dispersal. Journal of Insect Science, 10, e105.
Lindroth, C. H. (1974). Coleoptera, Carabidae. Handbooks for the identification of British insects. Royal Entomological Society, London. Volume 4(2).
Lindroth, C. H. (1986). The Carabidae (Coleoptera) of Fennoscandia and Denmark. Fauna Entomologica Scandinavica, 15(2), 233—497.
Luff, M. L. (1992). Provisional atlas of the ground beetles (Coleoptera, Carabidae) of Britain. Biological Records Centre, Huntingdon, England.
Luff, M. L., Eyre, M. D., & Rushton, S. P. (1989). Classification and ordination of habitats of ground beetles (Coleoptera, Carabidae) in North-East England. Journal of Biogeography, 16(2), 121.
Magura, T., Tothmeresz, B., & Molnar, T. (2008). A species-level comparison of occurrence patterns in carabids along an urbanisation gradient. Landscape and Urban Planning, 86(2), 134—140.
Matalin, A. V. (2006). Geographic variability of the life cycle in Pterostichus me-lanarius (Coleoptera, Carabidae). Entomological Review, 86(4), 409—422.
Molyneux, N., Soares, I., & Neto, F. (2013). WorldClim and DIVA software for modeling current and future climates at a 5 km resolution. Combating Climate Change, 2013, 325—342.
Musolin, D. L. (2007). Insects in a warmer world: Ecological, physiological and life-history responses of true bugs (Heteroptera) to climate change. Global Change Biology, 13(8), 1565—1585.
Nekuliseanu, Z. Z., & Matalin, A. V. (2000). A Catalogue of the ground-beetles of the Republic of Moldova (Insecta, Coleoptera: Carabidae). Pensoft, Sofia—Moscow.
Panagos, P., Ballabio, C., Meusburger, K., Spinoni, J., Alewell, C., & Borrelli, P. (2017). Towards estimates of future rainfall erosivity in Europe based on REDES and WorldClim datasets. Journal of Hydrology, 548, 251—262.
Poggio, L., Simonetti, E., & Gimona, A. (2018). Enhancing the WorldClim data set for national and regional applications. Science of the Total Environment, 625, 1628-1643.
Putchkov, A. (2011). Ground beetles of the Ukraine (Coleoptera, Carabidae). ZooKeys, 100, 503-515.
Putchkov, A. V. (2012). Faunisticheskij obzor karaboidnyh zhukov (Coleoptera, Caraboidea) Ukrainy [A review of the caraboids-beetles (Coleoptera, Cara-boidea) of Ukraine]. Ukrai'nskyj Entomologichnyj Zhurnal, 5, 3-44.
Putchkov, A. V. (2018). Zhuky-turuny (Coleoptera, Carabidae) transformovanykh tsenoziv Ukrajiny [Ground-beetles (Coleoptera, Carabidae) oftransformed cenoses ofUkraine]. I. I. Schmalhausen Institute ofZoology, Kyiv (in Ukrainian).
Schelvis, J., & Siepel, H. (1988). Larval food spectra of Pterostichus oblongopunctatus and P. rhaeticus in the field (Coleoptera: Carabidae). Entomologia Generalis, 13, 61-66.
Schwerk, A, & Jaskula, R (2018). Rare patterns of dorsal puncture in Pterostichus oblongopunctatus (Coleoptera: Carabidae). PeerJ, 6, e4657.
Sejnohova, H. (2006). The dynamics of carabid beetles (Carabidae) of floodplain forest in Southern Moravia. Acta Universitatis Agriculturae et Silviculturae Mendelianae Brunensis, 54(1), 107-120.
Sergeeva, T. K., & Grunthal, S. Y. (1988). Seasonal trophic dynamics ofPterosti-chus oblongopunctatus (Coleoptera, Carabidae). Zoologichesky Zhurnal, 67(4), 548-556.
Serrano, J. (2013). New catalogue ofthe family Carabidae ofthe Iberian Peninsula (Coleoptera). Ediciones Universidad de Murcia, Murcia, Spain.
Sharova, I. C. (1981). Zhiznennye formy zhuzhelic (Coleoptera, Carabidae) [Life forms of ground beetles (Coleoptera, Carabidae)]. Nauka Publishers, Moscow (in Russian).
Sharova, I. C., & Denisova M. I. (1997). Sezonnaya dinamika lesnykh populya-tsiy zhuzhelits roda Pterostichus (Coleoptera, Carabidae) [Seasonal dynamics of forest populations of ground beetles of the genus Pterostichus (Coleoptera, Carabidae)]. Zoologichesky Zhurnal, 76(4), 418-427 (in Russian).
Sharova, I. C., & Filippov, B. Y. (2003). Osobennosti zhiznennykh tsiklov zhu-zhelits (Coleoptera, Carabidae) v usloviyakh severnoy taygi [Peculiarities of life cycles of ground beetles (Coleoptera, Carabidae) in the conditions of Northern Taiga]. Zoologichesky Zhurnal, 82(2), 229-238 (in Russian).
Sharova, I. K., & Denisova, M. I. (1997). Seasonal dynamics of carabid populations from the genus Pterostichus (Coleoptera, Carabidae). Zoologichesky Zhurnal, 76(4), 418-427.
Siepel, H. (1988). A quantitative model on the population dynamics of Pterostic-hus oblongopunctatus F (Coleoptera, Carabidae) with special reference to the larval stage. Revue d Ecologie et de Biologie du Sol, 25(4), 435-450.
Sigida, S. I. (1993). Landshaftno-biotopicheskoe raspredelenie i jekologicheskaja harakteristika zhuzhelic (Coleoptera, Carabidae) Predkavkaz'ja i severnyh sklonov Central'nogo Kavkaza [Landscape and biotopic distribution and ecological characteristic of carabids (Coleoptera, Carabidae) of Ciscaucasia and northern slopes of the Central Caucasus]. Entomologicheskoe Obozre-nie, 72(1), 11-38 (in Russian).
Simon, E., Harangi, S., Baranyai, E., Braun, M., Fabian, I., Mizser, S., Nagy, L., & Tothmeresz, B. (2016). Distribution of toxic elements between biotic and abiotic components of terrestrial ecosystem along an urbanization gradient: Soil, leaf litter and ground beetles. Ecological Indicators, 60, 258-264.
Sklodowski, J. (2006). Anthropogenic transformation of ground beetle assemblages (Coleoptera: Carabidae) in Bialowieza Forest, Poland: From primeval forests to managed woodlands of various ages. Entomologica Fennica, 17, 296-314.
Sklodowski, J. (2014). Consequence of the transformation of a primeval forest into a managed forest for carabid beetles (Coleoptera: Carabidae) - a case study from Bialowieza (Poland). European Journal of Entomology, 111(5), 639-648.
Sklodowski, J., & Garbalinska, P. (2011). Ground beetle (Coleoptera, Carabidae) assemblages inhabiting Scots pine stands of Puszcza Piska Forest: Six-year responses to a tornado impact. ZooKeys, 100, 371-392.
Soboleva-Dokuchaeva, I. I. (1995). Osobennosti formirovanija fauny zhuzhelic (Coleoptera, Carabidae) agrocenozov Nechernozem'ja pri kontakte s lesom [Peculiarities of formation of ground-beetles fauna (Coleoptera, Carabidae) in agrocoenoses neighbouring to forests in the Non-Chernozem Area]. Ento-mologicheskoe Obozrenie, 74(3), 551-567 (in Russian).
Tamutis, V., Tamute, B., & Ferenca, R. (2011). A catalogue of Lithuanian beetles (Insecta, Coleoptera). ZooKeys, 121, 1-494.
Thiele, H. U. (1975). Interactions between photoperiodism and temperature with respect to the control of dormancy in the adult stage of Pterostichus oblongopunctatus F. (Col., Carabidae). Oecologia, 19(1), 39-47.
Thiele, H. U. (1977). Carabid beetles in their environments. Springer-Verlag, Berlin.
Thiele, H.-U., & Konen, H. (1975). Interaction between photoperiodism and temperature with respect to the control of dormancy in the adult stage of Pteros-tichus oblongopunctatus F. (Coleoptera, Carabidae). II. The development of the reproduction potential during the winter months in the field. Oecologia (Berlin), 19, 339-343.
Turin, H., Haeck, J., & Hengeveld, R. (1977). Atlas of the carabid beetles of the Netherlands. North-Holland Publishing Company, Amsterdam.
Ulrich, W., & Zalewski, M. (2006). Abundance and co-occurrence patterns of core and satellite species of ground beetles on small lake islands. Oikos, 114(2), 338-348.
Van Schaick Zillesen, P. G. (1985). Seasonal synchronization of the life-cycle of
Pterostichus oblongopunctatus (Coleoptera: Carabidae). Entomologia Gene-ralis, 11, 33-39.
Zalewski, M., & Ulrich, W. (2006). Dispersal as a key element of community structure: The case of ground beetles on lake islands. Diversity and Distributions, 12(6), 767-775. Zalewski, M., Sienkiewicz, P., Kujawa, K., Hajdamowicz, I., & Ulrich, W. (2012). Ground beetles on islands: On the effects of habitat and dispersal. Annales Zoologici Fennici, 49(3), 139-151.
Zamolodchikov, D. G. (2016). Prognoz rosta globalnoy temperatury v XXI veke na osnove prostoy statisticheskoy modeli [Global temperature rise forecast for the 21th century based on a simple statistical model]. Kompyuternyye Issledovaniya i Modelirovaniye, 8(2), 379-390.
Zamotajlov, A. S., & Nikitsky, N. B. (2010). Zhestkokrylye nasekomye (Insecta, Coleoptera) Respubliki Adygeja (annotirovannyj katalog vidov) [Coleopterous insects (Insecta, Coleoptera) of Republic of Adygheya (annotated catalogue of species)]. Adyghei State University Publishers, Maykop (in Russian).
Zherebcov, A. K. (2000). Opredelitel' zhuzhelic (Coleoptera, Carabidae) Respubliki Tatarstan [Handbook for the identification of ground beetles (Coleoptera, Carabidae) Republic of Tatarstan]. Kazan University Press, Kazan, Russia (in Russian).
