CC BY
17
УДК 595.762.12: 57.031
1T.R. Mukhametnabiev, 'RA. Sukhodolskaya, 2I.G.Vorobyova,
3A.L. Antsiferov, 4N.L.Ukhova
'Research Institute for Problems of Ecology and Mineral Wealth Use Tatarstan Academy of Sciences,
crtmatrix@gmail.com; ra5suh@rambler.ru 2Mariy State University, vigir@mail.ru 3Kostroma Museum of Nature, ancifer.ost@yandex.ru 4Visim State Nature Reserve, ukh08@yandex.ru
INFLUENCE OF GEOGRAPHIC LOCATION IN AREA AND DOMINANT FOREST FORMING SPECIES ON BODY SHAPE OF GROUND BEETLE PTEROSTICHUS OBLONGOPUNCTATUS FABRICIUS, 1787 (COLEOPTERA; CARABIDAE) IN TAIGA-BROADLEAF
GRADIENT IN RUSSIA
Pterostichus oblogopunctatus (Fabricius 1787) is a widespread carabid, that, despite of long lasted familiarity in Europe, is marginally studied in the rest of Palearctic. Latitude, longitude, height and other gradients are found there, yet transition with adjacent parts of Eastern Europe isn't steep. As a minor and come-at-able template of such environmental differences primeval protected areas of State Nature Reserve "Bolshaya Kokshaga", Volga-Kama State Nature Reserve and recovering after fire and felling respectively Visim Nature reserve and native fir forest in Kostrama oblast', Russia were accepted. Majority of samples demonstrated prevalence of P. oblongopunctatus and accounted for 2.142 individuals. Sampled material received morphometric measuring and its output was utilized for comparing samples of various environmental attributes in search of their impact on difference among groups of studied metric traits. Conducted research uncovered different contribution of sampling sites' geographic locality to animals of generally similar ecosystems. Moreover, more prominent input of different geographic locality is found to have remarkable, sometimes outright opposite effect in similar environment. Regardless of geographic location and dominating vegetation we recorded female-biased sexual size dimorphism in P. oblogopunctatus in relation to all measured traits.
Keywords: Carabidae; Pterostichus oblongopunctatus; geometric morphometrics; Permutational ANOVA; environmental impact.
Introduction
Carabidae as the second largest Coleoptera family which can be found inhabiting almost all terrestrial ecosystems. While diversity of the group is immense, majority of species is found to occupy niche of active soil surface predators and imbibe impact of environmental factors of a vast part of inhabited area. Distance from beginning of food chain and soil dwelling of larvae make animals susceptible to prolonged environmental effect exposition even among young specimen, that makes them good object of bioindication studies, especially in case of abundant species (Kotze et al., 2011). Explicit monophyletic taxonomy of the group compared to other terrestrial Coleopteras reasons a distinct body composition, what simplifies geometric morphometric approach, where multiple traits, describing body shape are considered (Magura et al., 2006). Considering that, impact of environmental factors could manifest as difference in traits of body shape among animals of different localities and corresponding environmental
impact (Baranovska, Knapp, 2013).
Pterostichus oblongopunctatus (Fabricius, 1787) is a small-medium size widespread carabid, that is omnipresent throughout majority of European ecosystems (Brygadyrenko, 2016) with higher occurrence in litter-rich soils of humid deciduous broadleaved forests (Irmier 2007; Magura et al., 2001). Despite of that, the species is noted in habitats of significant temperature span and found susceptible to multitude of following environmental factors (Avtaeva et al., 2019). Research of sensitivity of animal's metric traits to both human activity (Lagisz, 2008) and natural conditions are noted (Belskaya et al., 2019). The animal is mobile and notable quantity of specimen is found in ordinary pitfall traps, hereby providing raw material for statistical analysis. Despite of textbook attribution to ecosystems of Europe (Kryzhanovsky, 1965), given species is found abundant further in Palearctic. Conducted research provides insight on their presence and abundance in Mesouralic and Siberian ecosystems (Sukhodolskaya
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Fig. 1. Locality of researched territories on map of Europe (Source image is accepted on terms of CC BY-SA 3.0 (Europelaea location map.sv, 2012)
et al., 2018). All given geographic areas of research are ecosystems with homogenous arboreal composition, characterized by a single dominant species even in non-climax ecosystems and cover large geographic span. Influence of different ecosystems, geographic location and climate is noted for similar species (Sukhodolskaya, Ananina, 2017), yet remain not entirely researched in huge span of environmental factors for P. oblongopunctatus.
Considering that, following geographic localities provide a minor model of researched ecological influence on the species.
Area of Volga-Kama State Nature Biosphere Reserve encompasses two partition with different environmental conditions, which were regarded as different geographic localities. The reserve is an object of the annual sampling as a collaboration agreement between administration of the Reserve and Laboratory of Biomonitoring of The Institute of Problems in Ecology and Mineral Wealth Use Tatarstan Academy of Sciences. Macrofauna samples were utilized for description of taxonomic diversity whereas Carabid animals were used for the following research.
Territory of State Nature Reserve «Bolshaya Kokshaga» is situated in basin of Bolshaya Kokshaga river. The reserve is majorly covered by pine forests and geographically situated closely to Volga-Kama reserve.
Visimskiy State Nature Reserve is characterized by recovering from several vast wildfires dating 1920, 1998, 2010 in fir dominated areas. Kalinki
village, Kostroma oblast' also represents fir dominated ecosystem recovery after felling of the native fir forest in recent decades.
Last geographic localities received separate treatment due to lack of overlapping values of major forest forming specie and condition of active recovery.
Body shape morphology is one of primary and utmost simple ways of collecting characteristics of an individual animal and it's applicable for large samples. While complexity of each body part is immense, the utilized methodology in reason of its age prioritized simplicity and robustness. Thus, while certain details might remain unknown, simple metrics should provide insight on most prominent traits. Hereby, we used data on beetles' measurements, which had earlier been obtained using Sukhodolskaya's method (Sukhodolskaya, Saveliev, 2016). Considering the described flaws, raw metric data was accepted as is and measuring method was regarded as an initial condition rather than researcher's choice.
Throughout 4 years of research in Volga-Kama reserve, the given species was selected as the most abundant carabid species and is widespread in overwhelming majority of samples independently of values of sample's attributes.
Conducted research was initially purposed to perceive difference of body shape among sampled Carabids and to reveal influence of environmental factors on studied multitude of metric traits. Each animal was regarded as an object and its independent attributes comprised sample specific values of environmental factors' attribute. These values, except for animal specific sex attribute, are shared among the entire sample. Each of metric traits was regarded as a dependent attribute. Values of these are unique for each specimen.
Thus, the research was required to demonstrate difference among groups of dependent values, which are selected according to associated independent values.
Material and methods
Sampling area is presented on the map (fig. 1).
Volga-Kama State Nature Biosphere Reserve is a protected area near Kuybyshev reservoir, consists of two partitions:
1) Raifski site is situated on southern border of taiga ecological zone. Majority of its area is covered by coniferous forests (Pinus sylvestris as major
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Table 1. Listed values of analyzed attributes (samples, further disregarded during an attribute specific analysis are given in
italic)
Attribute Value Animal quantity
Birch 148
Major forest forming Linden 921
species Pine 359
Fir 714
Raifski 587
Saralinski 686
Locality Bolshaya Kokshaga 155
Visimsky 368
Kalinki 346
18 18
24 245
30 88
Sampling site (Saralinsky partition) 38 34
40 2
42 1
54 176
55 97
56 25
25 43
33 58
37 43
47 11
Sampling site (Raifsky partition) 58 33
63 41
80 118
81 78
84 84
86 78
Sampling site (BK) 90-91 155
PZP-2 96
Sampling site (Visimski) PZP-7 118
PZP-19 90
PZP-20 64
Sampling site (Kalinki) none 346
Female 996
Sex Male 1138
Unspecified 8
forest forming tree species), southern parts of area are covered with deciduous broadleaved forests, (majorly composed of Tylia cordata). The partition is also characterized by higher perineal precipitation. Despite of an elevation span of 86-140 m above sea level, it's located slightly lower than surrounding area,
thus ground water is abundant (Gasanov et al., 2014; Burganova et al., 2014).
2) Saralinski site is situated along the coast of Kuybyshev reservoir. Oppositely to Raifski site, majority of area is covered by linden forests, yet significant amount of pine forests is found along the coast (major forest forming species are the same) (Shaykhutdinova et al., 2014).
Despite of both partitions having ordinal notation of quarters, no sampling was done in quarters with similar index. Thus, no sample required referring, other than its number, whereas partition affiliation is given in Table 1 alongside with other attributes.
Animals from Bolshaya Kokshaga reserve were sampled in a single quarter №90-91 due to proximity to of the stationary observation point. The area is covered with pine dominated forests and represents similarities with those of Volga-Kama reserve.
Visimskiy reserve is represented by the set of soil zoologic plots (rus. Pochvenno-Zoologicheskaya Ploshchadka - PZP) (Sukhodolskaya et al., 2018) with each of them corresponding to a different condition of fir forest recovery. Site PZP19 represents primeval fir forest with the utmost density of undergrowth and presumed to be the most favorable for Carabids due to that and the following heat accumulation. PZP7 was affected by fire of 1920 and underwent introduction of significant amount of deciduous species syne, which currently are replaced by fir. PZP20 corresponds to fires of both 1998 and 2010. Due to weak recovery after the first fire, not much combustive biomass was retained, resulting in lesser damage of the second one. PZP2, while affected only by fire of 2010, exhibits the lowest recovery and, oppositely to PZP19, assumed as the least favorable environment for Carabids.
Animals from Kalinki were collected in area of a single forest felling, that was referred as a solitary sampling site, similarly to sample № 90-91 of BK reserve. In Volga-Kama reserve in period of 2014-2017 totally 70 samples were taken. Gathered samples comprise overall exposition of 3562 days per trap. In Bolshaya Kokshaga reserve animals were sampled in 2014. Beetles from Visimskiy reserve and Kalinki were sampled during period of 2014 and 2015
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10 mm scale terminus 1 (index order is .irrelevant)
10 mm scale
terminus 2
Fig. 2. Placement of morphometric measurements: A - Elytra length as distance between posterior end ofscutellum and terminus of right elytron (in
absence case of intact right elytron, left one is acceptable); B - Elytra width as distance between
anterior-distal corners of elytra; V - Pronotum length measured along of central furrow pronotum; G - Pronotum width as distance between posterior corners of pronotum; D - Head length as distance
between labrum and juncture of occiput and postgena. (This trait despite of its clear wording, is prone to erroneous deviations due to mobility of head joints and following inconsistency of the anterior terminus); E - Head width as distance between proximal innermost sides of eyes.
accordingly.
Studied animals were captured using pitfall traps with saline solution. Each sampling site comprised a direct line of traps with over 10 m distance betwixt each. Sampling trap exposition lasted approximately 5 days length (Barber, 1931). Subsequently, animals were preserved in 70% alcohol prior to taxonomic differentiation (Sasakawa, 2007) and were put straightened on cotton paddings thereafter.
Among them, 2142 specimens were accepted for following measuring. Subsequently, according to sampling data, following values of independent attributes are initially given for animals of each sample (tab. 1).
Utilized morphometric data corresponded to six linear scalar float recordings as dependent variables and is given for following values (denoting letters are given as Cyrillic transliteration of reference material's source) by R.A. Sukhodolskaya (Sukhodolskaya, Saveliev, 2016) (fig. 2).
Morphometric data was collected from images taken by Nikon D5100 camera with custom opaque
light disperser and a box with opaque reflective surface. Measurements were made using program, designated specifically for the given method of measurement and utilized distance between manually pointed out elements of photos' arrays as terminal point of measurements and fiducial scale, using the last to bind real scale to array output data.
According to distribution of six dependent parameters of given attributes by values of categorical independent variables, 432 groups of Volga-Kama and BK reserves and 60 groups of Fir dominated areas with quantity equal or more than 3 specimens, what is minimum for Shapiro-Wilks test, as the most powerful (Shapiro, 1965; Yap6 Sim, 2011; Jones et al., 2001), were found and analyzed.
While all test W values were found to range between 0.998-0.666, p-value proved to be less than 0.05 for 94 samples of VK and BK reserves and 37 in fir dominated localities. This considering, that groups of different dependent attributes and the same value of independent were not always entirely parametric or not, made usage of parametric multivariate analysis nigh impossible.
Thus, initial overview was performed using non-parametric 2-way PERMANOVA. The analysis was conducted using PAST 3 with default permutation kwarg set to 9.999 (Hammer, 1999; Anderson 2001). Explicit difference among samples and interaction were handled visually («Lecture 27 STAT 512», 2011; Hunter, 2007; Müllner, 2001).
Unfortunately, method specific proportions of the metric traits as well as lack of normality set PCA impractical (Jolliffe, Cadima, 2016) and confined the research to mere visual handling with reliance on standard deviation.
Results
Significant reciprocal interaction (tab. 2) was found among following grouping attributes.
Visual depiction of found interactions is given on the following plots for each pair of interacting attributes. All metric traits of approximately similar values are grouped by subplots and each of them is marked with subplot specific greyscale marking. In case of each metric trait means for each attribute 1 categories of each of attribute 2 categories samples and were plotted with scalar value annotations given on y axis. Hereby we revealed the increase of animals' matric traits in birch dominated habitats and decrease of it in case of pine in Raifski partition and the opposite effect in Saralinki partition (fig. 3).
This is the most remarkable demonstration of differences among metric traits.
Influence of sexual dimorphism, manifested as larger females, is notable, yet the interaction is
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Table 2. PERMANOVA output for given pairs of attributes in Volga-Kama and Bolshaya Kokshaga reserves
Pairs Source Sum of sqrs df Mean square F p
Reserve partition and major forest forming species Ecosystem 52.83 2 26.415 45.134 0.0001
Area 122.23 2 61.117 104.43 0.0001
Interaction -423.74 4 -105.94 -181.01 0.0121
Residual 831.07 1420 0.585
Total 582.39 1428
Sex and major forest forming species Ecosystem 52.83 2 26.415 43.929 0.0001
Sex 31.07 2 15.535 25.835 0.0001
Interaction -355.36 4 -88.84 -147.75 0.0001
Residual 853.85 1420 0.601
Total 582.39 1428
Reserve partition and sex Area 122.23 2 61.117 137.2 0.0001
Sex 31.07 2 15.535 34.874 0.0001
Interaction -203.47 4 -50.867 -114.19 0.0001
Residual 632.56 1420 0.445
Total 582.39 1428
obscure (fig. 4, 5).
For the only remaining non-interacting categorical attribute of Quarter one-way PERMANOVA provides following results in table 3. The square matrix output was calculated for Euclidean distance scalars and was utilized for further dendrogram visualization (fig. 6). Visual depiction of difference among samples is given on figure 7.
Two-way PERMANOVA of geographic locality and sampling site detected no interaction among any of attributes' values in samples of Visimskiy reserve
and Kalinki sample (tab. 4). One-way analysis, while separating animals from Visim Reserve according to sampling site variable, regarded animals from Kalinki as a sample of a single sampling site, thus effectively eliminating need of considering geographic locality separately (fig. 8, 9).
Hereby, prominent and clear interaction of forest forming species and geographic locality is noted.
Discussion
By all means, body shape of any animal is influenced
Fig. 3. Plot visualization of interaction between attributes of geographic localities and all major forest forming species. a) Means of elytra lengths; b) Means of elytra width (B), pronotum length (V), pronotum
width (G); c) Means of head length (D), head width (E)
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Fig. 4. Plot visualization of interaction between attributes of major forest forming species and extent of sexual dimorphism among sampled animals. a) Means of elytra lengths; b) Means of elytra width (B), pronotum length (V), pronotum width (G); c) Means of head length (D), head width (E)
6.10-
3.0
201.91.81.71.6 1.5 -1.41.3
Raifski
Saralinski BK
Raifski
Saralinski BK
Raifski
Saralinski
a) — (a
m,A Ь)
— ЕВ fv ■■■ f G - ID IE
ni,B m, V tllG С) m,D tll,E
Fig. 5. Plot visualization of interaction between attributes of sexual dimorphism and geographic localities. a) Means of elytra lengths; b) Means of elytra width (B), pronotum length (V), pronotum width (G); c) Means
of head length (D), head width (E)
by environmental factors, be these favorable or not. Auspicious factors might not always be the reason for increase of body size and provide other benefits like reproduction and mobility (Muona, Rutanen, 1994), yet are most likely to manifest in size (Lagisz, 2008). Hereby that is presumed, but the emphasis is made on highlighting difference among sets of scalar values.
PERMANOVA of categorical attributes revealed explicit simple interaction among multitude of them, which received visual treatment.
Dependence of metric traits on sexual dimorphism is apparent among multitude of Carabids (Sukhodolskaya et al., 2016), including similar species and affects all explored metric traits (Brygadyrenko, Korolev, 2015). Studied P. oblongopunctatus is no different, for which expectedly larger females were noted in our research. Visual depiction of interaction among groups of sex attribute and other participant traits demonstrated weak and obscure outcome
compare with sampling locality/major forest forming pair. Thus, no novelty is revealed for interactions of sex attributes and the emphasis was made on the remaining pair.
Received plots for pair of major forest forming species and reserve partition demonstrate relatively uniform influence of interaction on all researched metric traits. Means of almost each trait among animals from Raifski partition have minimal value in samples from pine ecosystems, intermediate in
Table 3. One-way PERMANOVA of Sampling site's quarter attribute in Volga-Kama and Bolshaya Kokshaga reserves
in
Permutation N 9999
Total sum of squares 582.4
Within-group sum of squares 437.7
F 24.52
p (same) 0.0001
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Fig. 6. Tree of Euclidean distances of PERMANOVA as dissimilarity among samples of Volga-Kama and Bolshaya Kokshaga reserves' quarters. Body shape
difference among two major clusters of animals from quarters 55, 25, 63 and the rest of samples is demonstrated
Fig. 7. Visualization of distribution of metric traits' values from samples of animals from Volga-Kama and
Bolshaya Kokshaga reserves. «Grey branch» box is used for representing quarters of conforming color annotation on figure 6 for sake of brevity. A - variation in elytra lengths, B - variation in elytra width, V - variation in pronotum length, G - variation in pronotum width, D - variation in head length, E -variation in head width. o - outliers, ▲ - means, — - medians, — -1.5 intequartile interval whiskers, □ - II-III quartile span boxes
linden and achieves maximum relatively birch. Exactly opposite results are found for animals of Saralinski partition. Values of head length attribute are not entirely consistent relatively the flora, but influence of partitions depicted as the slope is not different to other traits.
Even more prominent difference of animals from not very distant BK reserve is noted as well. While distinct reasons of this might retain obscurity, the emphasis could be made on geographic locality as the major factor.
The dendrogram of pairwise distance calculation demonstrated body shape difference among two major clusters of animals from quarters 55, 25, 63 and the rest of samples. Also, despite of absence of significant interaction with other independent values, affiliation of similar (distance 0.332, with p 0.724) samples from quarters 25 and 63 with Raifski partition and pine forests is noted. Regarding the 90-91 samples of BK, considering prominent difference, the lesser accounting could be given to trait A and B, whereas values of the rest of them are found much smaller. Visual handling of distribution plots revealed animals from the remaining majority of samples being generally larger than in any of the «outlined branch».
One-way per sampling site analysis and locality analysis of animal from recovering fir dominated territories revealed drastic difference of metric structure of larger animals from Kalinki, yet the difference is proved to be significant among each of samples too. Despite of pertaining to different stages of recovery, overall difference among animals from Visim reserve was much less in compare to animals from Kalinki of Kostroma oblast'. This can be considered as an evidence of geographic locality's impact, yet this also does eloquently represent encompassing multitude of unknown factors by a single attribute value due to lack of felling sites in Visim reserve. This could also be asserted by a significant difference between animas from primeval site (plot 19) of Visimski reserve and BK with Volga-Kama reserves.
In accordance to presumption of the most favorable environment in Visim reserve, larger animals indeed were found in partition with primeval ecosystem, yet not much difference is seen among the remaining samples from the recovering sites. Animals from fire affected sampling sites demonstrate more similarities, yet difference of metric traits' values and their distribution is just less prominent. This could be generalized as overally more notable susceptibility to disturbance of ecosystems, rather than any specific factor of disturbance in given range.
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Table 4. Results of one-way PERMANOVA of sampling site's quarter attribute in fir dominated territories
Fig. 9. Visualization of distribution of metric traits' values from samples of animals from fir dominated ecosystems, demonstrating difference among the metric traits of animals from Kalinki and different
clusters of Visim reserve. A - variation in elytra lengths, B - variation in elytra width, V - variation
in pronotum length, G - variation in pronotum width, D - variation in head length, E - variation in head width. o - outliers, ▲ - means, — - medians, — -1.5 intequartile interval whiskers, □ - II-III quartile span boxes
Permutation N 9999
Total sum of squares 516.5
Within-group sum of squares 342.5
F 90.04
p (same) 0.0001
Fig. 8. Tree of Euclidean distances of PERMANOVA as dissimilarity among samples from Fir dominated localities with distinguished difference of animals from Kalinki site and lesser extent of it at all sampling sites of Visim reserve
This could fit observations of similar impact of other different conditions as well (Zygmunt et al., 2006). Although quantity of samples wasn't given, lesser difference between sites of different environmental disturbance might indicate, that susceptibility of P. oblongopunctatus as a generalist species to fire is not strong and might even be argued as neutral rather than negative if quantity is prioritized over individuals' size (Muona, Rutanen, 1994).
Conclusion
A range of factors, including spatial ecological gradients, historical climate changes, and physical barriers to gene flow might have contributed to the organization of body size variation in carabids. Our work is the first attempt to describe carabid beetle morphometric variation on the basis of several traits analysis which we have further improved by several populations research sampled in different regions. We have demonstrated that an informative approach ought to take into account geographical position and environmental impact on body size variation. A combination of intraspecies and functional approaches provides extra information confirming that the analysis of species' morphofunctional features is a promising and common approach to evaluate functional diversity. Our work suggests that a functional approach can reveal the interactions between specific locality and the factors affecting forest ecosystems, not only spatially but also temporally.
Future research should be focused on the role of environmental factors combined study when investigating body size variation in the certain species of Ground Beetles. Moreover, as we have found that several factors affect not to the same degree the studied species variation, the complexity of the forest ecosystems must be considered when planning to use carabids for providing ecosystem services.
Acknowledgements
We would like to express gratitude to all employees of Volga-Kama reserve for the given possibility of collaboration and Dmitry Vavilov from biomonitoring
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laboratory of The Institute of Problems in Ecology and Mineral Wealth Use for assistance in taking animals' photos.
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32. Zygmunt P.M.S., Maryanski M., Laskowski R. Body mass and caloric value of the ground beetle (Pterostichus oblongopunctatus) (Coleoptera, Carabidae) along a gradient of heavy metal pollution // Environmental Toxicology and Chemistry. 2006. №25(10). P. 2709-2714.
Мухаметнабиев Т.Р., Суходольская Р.А., Воробьева И.Г., Анциферов А.Л., Ухова Н. Л. Влияние географического положения в ареале и доминирующей растительности на изменчивость размеров тела жужелицы Pterostichus oblongopunctatus fabricius, 1787 (Coleoptera; Carabidae) в таежных и широколиственных
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лесах России.
Pterostichus oblogopunctatus ^аЬйс^, 1787) -распространенная жужелица, но, несмотря на хорошую изученность в Европе, мало исследована в остальной Палеарктике, где наблюдаются долготная и широтная зональности, но градиент с прилегающими частями Восточной Европы не резкий. Как доступный шаблон подобной разницы среды в качестве исследуемых были взяты защищенные территории Государственного природного заповедника «Большая Кокшага», Волжско-Камского государственного природного биосферного заповедника и восстанавливающиеся естественные еловые леса в Висимском природном заповеднике (после пожара) и в Костромской области (после рубки). Большинство проб продемонстрировали преобладание Р. oblongopunctatus, насчитывавшего 2142 экземпляра. Проведен морфометрический
анализ по шести мерным признакам. Показано, что основным фактором, влияющим на морфо-метрическую изменчивость жужелицы исследуемого вида, является регион обитания, поскольку даже в сходных, но расположенных в разных регионах экосистемах изменчивость размеров жуков различна. Более того, она может быть разнонаправленной в однотипных биотопах в зависимости от региона обитания. Независимо от места обитания и доминирующей растительности у Р. oblogopunctatus зарегистрирован половой диморфизм по всем исследованным признакам - самки больше самцов.
Ключевые слова: Carabidae; Pterostichus oblongopunctatus; геометрическая морфометрия; пермутационный анализ вариации; экологические факторы.
Информация об авторах
Мухаметнабиев Тимур Ринатович, младший научный сотрудник, Институт проблем экологии и недропользования АН РТ, 420087, Россия, г. Казань, ул. Даурская, 28, E-mail: crtmatrix@gmail.com.
Суходольская Раиса Анатольевна, кандидат биологических наук, старший научный сотрудник, Институт проблем экологии и недропользования АН РТ, 420087, Россия, г. Казань, ул. Даурская, 28, E-mail: ra5suh@rambler.ru.
Воробьева Ираида Геннадьевна, кандидат биологических наук, доцент, Марийский государственный университет, 420039, Россия, г Йошкар-Ола, ул. Школьная, 57, E-mail: vigir@mail.ru.
Анциферов Анатолий Леонидович, кандидат биологических наук, старший научный сотрудник, Костромской государственный историко-архитектурный и художественный музей-заповедник, 156000, Россия, г. Кострома, пр. Мира, 7, E-mail: ancifer.ost@yandex.ru.
Ухова Надежда Леонидовна старший научный сотрудник, Висимский государственный природный биосферный заповедник, 624144, Россия, Свердловская область, г. Кировград, ул. Ст. Разина, 23, E-mail: ukh08@yandex.ru.
Information about authors
Timur R. Mukhametnabiev, Junior Researcher, Research Institute for Problems of Ecology and Mineral Wealth Use of Tatarstan Academy of Sciences, 28, Daurskaya st., Kazan, 420087, Russia, E-mail: crtmatrix@gmail.com.
Raisa A. Sukhodolskaya, Ph.D., Senior Researcher, Research Institute for Problems of Ecology and Mineral Wealth Use of Tatarstan Academy of Sciences, 28, Daurskaya st., Kazan, 420087, Russia, E-mail: ra5suh@rambler.ru.
Iraida G. Vorob'eva, Ph.D., Assistant Professor, Mariy State University, 57, Shkolnaya st., Yoshkar Ola, 420039, Russia, E-mail: vigir@mail.ru.
Anatoly L. Antsiferov, Ph.D., Senior Researcher, Kostromа State Historic, Architectural and Art Museum-Reserve, 7, Mir Av., Kostroma, 156000, Russia, E-mail: ancifer.ost@yandex.ru.
Nadeyhda L. Ukhova, Senior Researcher, Visimskiy Nature Reserve, 23, St. Razin st., Kirovgrad, 624144, Russia, E-mail: ukh08@ yandex.ru.
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российский журнал прикладной экологии
