Bull 721 Bull. Iraq nat. Hist. Mus. (2025) 18 (3): 721-738. https://doi.org/10.26842/binhm.7.2025.18.3.0721 ORIGINAL ARTICLE LANDSCAPE-DEPENDENT VARIABILITY OF MORPHOMETRIC TRAITS IN CARABUS GRANULATUS L., 1758 (COLEOPTERA, CARABIDAE) Roman Gorbunov*,**♦, Viktor Aleksanov***, Rifgat Shagidullin*, Sergey Berdnik* and Raisa Sukhodolskaya*,****,***** * Institute for Problems of Ecology and Mineral Wealth Use of theTatarstan Academy of Sciences, Russian Federation, 420087, Kazan. ** Belogorye State Nature Reserve, Russian Federation, 309342, Borisovka settl. ***State Budgetary Institution of Kaluga Region “Park Directorate”, Russian Federation, 248000, Kaluga. **** Kazan State Medical University, Russian Federation, 420012, Kazan. ***** Volga-Kama State Natural Biosphere Reserve, Russian Federation, 422623, Sadovyi settl. Corresponding author: mailto:xobglor@gmail.com Received: 28 Feb. 2024, Revised: 24 May 2025, Accepted: 29 May 2025, Published:20 June 2025 This work is licensed under a Creative Commons Attribution 4.0 International License ABSTRACT We examined morphological variations in Carabus granulatus L., 1758 (Coleoptera, Carabidae) populations across different habitats in the Kaluga Region. Discriminant analysis revealed subtle but significant morphological differences between populations, with pronotum length being the key discriminating trait. Morphological variability was primarily driven by historical microevolutionary processes rather than direct phenotypic adaptation to specific habitats. Local ecological factors, such as microclimate and resource availability, had a greater influence on morphometric traits than the general habitat type. Populations in the eastern part of the Kaluga Region exhibited smaller body sizes due to intensive forest use. Fragmented forests displayed notable differences in morphometric traits, likely resulting from founder effects. Beetles from various localities showed distinct patterns: populations in the Oka River valley populations were smaller, while park habitats exhibited increased variability. Populations in Kaluga occupied an intermediate position between wild and disturbed forests. The study emphasizes the importance of considering both phylogenetic and ecological factors in understanding morphological differences in ground beetles, highlighting the complex interplay between historical, environmental, and anthropogenic factors shaping population- level morphological variation in C. granulatus. Keywords: Biogeoregion, Carabus, Discriminant analysis, Ground beetles, Morphometry. BULLETIN OF THE IRAQ NATURAL HISTORYMUSEUM Iraq Natural History Research Center & Museum, University of Baghdad https://jnhm.uobaghdad.edu.iq/index.php/BINHM/Home Copyright © Bulletin of the Iraq Natural History Museum Online ISSN: 2311-9799, Print ISSN: 1017-8678 https://doi.org/10.26842/binhm.7.2025.18.3.0721 https://orcid.org/0009-0006-3849-2006 https://orcid.org/0000-0002-4584-8457 https://orcid.org/0000-0002-3321-0923 https://orcid.org/0009-0003-8261-2421 https://orcid.org/0000-0002-6402-7152 mailto:xobglor@gmail.com https://creativecommons.org/licenses/by/4.0/ https://jnhm.uobaghdad.edu.iq/index.php/BINHM/Home 722 Bull. Iraq nat. Hist. Mus. 18 (3): 721-738. Landscape-dependent variability INTRODUCTION Studying how organisms respond to ecological factors is essential for developing biomonitoring models and for gaining insights into the phylogenetic pathways of taxa. It is well-established that animals exhibit dynamic adaptive and microevolutionary changes in response to environmental alterations (Newman, 1995; Staton et al., 2001; Hauser et al., 2003). These transformations can occur independently or in conjunction, forming complex relationships in which the local environment influences their expression (Peiman and Robinson, 2017). Species-specific traits can be viewed as components of biodiversity, affecting how organisms react to disturbances and shifting environmental conditions, with implications at both population and broader ecological levels (Lavorel and Garnier, 2002). Therefore, selecting particular traits for study is crucial for determining the ecological data obtained. Body size is a fundamental characteristic of organisms, varying with environmental conditions and subject to selective pressures. In insects, body size is linked to numerous life- history traits, including reproductive rate, dispersal capability, duration of ontogeny, competition, and the number and size of offspring. These factors significantly shape the interactions between insects and their environment (Stearns, 1998). Current research indicates that the sizes and shapes of individual beetle organs vary across habitats with different vegetation types and levels of anthropogenic impact (Luzyanin, 2023; Khomitskiy et al, 2024). For instance, studies on the ground beetle Poecilus cupreus (L., 1758) have demonstrated that its body size diminishes in agrocenoses, with both the agrocenosis conditions and the type of vegetation significantly affecting the beetles morphology (Sukhodolskaya et al., 2017). C. granulatus is a transpalaearctic, and hygrophilous eurytopic species with a one-year life cycle. It is a spring breeder species without larval diapause, but with obligatory diapause in adult individuals. Both adults and larvae are predators (Turin et al., 2003). The objective of this study was to identify (i) the differences among Carabus granulatus L., 1758 individuals from various localities, (ii) the variations between habitat types within the Kaluga region or specific sites within a given locality, and (iii) how habitat type (defined by dominant tree species) and geomorphological zonation impact these differences. MATERIAL AND METHODS Characteristics of the study area: The Kaluga Region is located in the center of the East European Plain, south of Moscow, primarily within the upper basin of the Oka River – the largest right tributary of the Volga. Its northern and western parts belong to the zone of coniferous-broadleaved forests (Lavrenko, 1947), which is classified in Europe as the Boreal biogeoregion (European Environment Agency, 2016). The zonal vegetation is represented by spruce-broadleaf and broadleaf-spruce forests, dominated by European spruce Picea abies (L.) H. Karst., 1881, small-leaved linden Tilia cordata Mill, 1768, and pedunculate oak Quercus 723 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Gorbunov et al. robur L., 1753. Swamps are common. The relief was formed as a result of the Moscow glaciation (Esipov, 2005). The southern and eastern parts of the Kaluga region belong to the zone of broadleaf forests, also known as the Continental biogeoregion. This area has been influenced by the older Dnieper glaciation and is more distinctly dissected by a network of rivers, with remnants of pre-Quaternary relief preserved in its eastern part. Due to prolonged economic use, the eastern part of the Kaluga region is practically devoid of minimally disturbed forests, while the southern part retains small areas of undisturbed broadleaf forests within the protective belt that defended the southern borders of the Moscow state, where European ash Fraxinus excelsior L., 1753 is present in the stand (Bobrovsky and Abatises, 2002). Characteristics of the sampling sites: The material for this study consisted of specimens of C. granulatus collected from 12 sampling sites in 10 localities (Tab. 1.) between 1996 and 2021. A locality is defined as a group of sampling sites located within 5 km of each other and not separated by significant barriers (Map 1). Geographic coordinates are provided in the order of latitude – longitude. High intensity resource use refers to non-forest use of the territory in the recent past or to a significant frequency and intensity of logging. This is identified by: 1) historical maps and satellite images, 2) the current structure of the stand, characterized by young trees or older ones that are regularly planted, with reduced age diversity of trees. Low intensity use: the opposite condition of the same signs, as well as the presence of plant species sensitive to anthropogenic pressure. Medium intensity use: in the recent past, the forest was not completely logged, but deadwood was collected, and there may have been livestock grazing, etc. Forest arrays are classified as follows: small — up to 1.5 sq. km, medium — 10-20 sq. km, large — more than 100 sq. km. Areas with a high perimeter-to- area ratio are also classified as small. Table (1): Locations of carabid beetle trapping, grouped be vegetation zone and geomorphological zones [by Esipov (2005)]. Locality Type of habitat (by groups of dominant trees) Position in the landscape Dominant plant species Size of forest array Intensity of resource use over 100 years Broadleaved forests vegetation zone Moraine-glacial gently undulating plain of the Dnieper Glaciation Kireykovo (Kir)* 53.6252 35.8647 Broadleaved forests Watershed Q. robur F. excelsior T. cordata Aegopodium podagraria L. Allium ursinum L. Large Low Moraine gently sloping plain of the Dnieper Glaciation with inherited post-Quaternary relief Bragino (Brg) See text for details Small- and broadleaved forest Big River Valley See text for details Small High Gremyachevo (Gr) 54.2373 36.2588 Small-leaved forest Big River Valley Alnus glutinosa (L.) Urtica dioica L. Small High 724 Bull. Iraq nat. Hist. Mus. 18 (3): 721-738. Landscape-dependent variability Tarusa (Tar) 54.7257 37.1989 Small-leaved forest Big River Valley Salix sp. Alnus incana (L.) Moench U. dioica Small High Trubetskoe (Trb) 54.6419 37.1597 Broadleaved forests Watershed T. cordata A. podagraria Carex pilosa Small High Sivtsevo (Siv) 54.6316 37.0491 Broadleaved forests Watershed T. cordata A. podagraria Mercurialis perennis L. Small High Mixed forests vegetation zone Moraine gently undulating plain of the Moscow Glaciation Kaluga, Bor (K-Br) 54.5436 36.1955 Spruce forest Watershed Picea abietis Pinus sylvestris Betula pubescens Galeobdolon luteum Huds. Oxalis acetosella L. Medium Medium Kaluga, Ovrag Mozhayka (K-Mz) 54.4876, 36.2402 Small-leaved forest Small River Valley Betula pendula P. sylvestris Salix sp. Á. Incána Corylus avellana L. Small Medium Flat glacial plain of the Moscow Glaciation Milyatino (Mil) 54.4811 34.3551 Spruce forest Watershed P. abietis A. glutinosa Populus tremula L. Padus avium C. pilosa Geum rivale L. Medium Medium Budnyan-skiy (Bud) 54.2749 33.9226 Spruce forest Watershed P. abietis P. tremula C. avellana C. pilosa G. luteum M. perennis Small Low * – Tables and images below use these letter abbreviations. The locality “Bragino” includes three habitats: 1. Floodplain of the Oka River, willow-gray alder thicket with Urtica dioica, 54.4193 36.6749. 2. Valley of a small river (first-order tributary of the Oka), gray alder-thicket with Padus aviumMill., 54.4108 36.6678. 3. Gully with a stream (tributary of the small river), linden thicket with Corylus avellana and Carex pilosa Scop., 54.4044 36.6702. 725 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Gorbunov et al. Map (1): Kaluga region, studied localities. Research technique: The following morphological traits of the beetles were measured: length and width of the elytra, length and width of the pronotum, head length, and distance between the eyes. A total of 4,252 specimens were measured. After sampling the beetles and determining their species and sex, all individuals were photographed, and their parameters were measured using a custom program written in Python 2.7, utilizing the Num Py and Open CV libraries. (Mukhametnabiev, 2018). The scheme for measuring the beetles is presented in Plate (1). Linear discriminant analysis was performed by STATISTICA 10 software. 726 Bull. Iraq nat. Hist. Mus. 18 (3): 721-738. Landscape-dependent variability Plate (1): Illustration of measurements: EL – elytra length, EW – elytra width, PL – pronotum length, PW – pronotum width, HL – head length, DE – distance between the eyes. 727 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Gorbunov et al. RESULTS Discriminant analysis by locality According to the results, the populations of C. granulatus are well-discriminated and differ significantly (Wilks' Lambda: 0.41 approx. F (54.224) = 75.96; p<0.001). The coefficients of classification functions show that the length of the pronotum is the most important trait for discrimination. Other characteristics, with the exception of pronotum width, have a smaller and approximately equal contribution. The contribution of pronotum width to discrimination is extremely low. Squared Mahalanobis Distances (Tab. 2.) vary from 0.20 to 14.70. The largest distances are observed between the pairs Milyatino-Sivtsevo (14.70) and Milyatino-Gremyachevo (12.24). The smallest distances between the values belong to Tarusa and Bragino (0.20) and Tarusa- Trubetskoy (0.22). Table (2): Squared Mahalanobis distances (locality), p<0.001 Kir Br Gr Tar Trb Siv K-Br K-Mz Mil Bud Kir - Br 3.41 - Gr 4.42 0.53 - Tar 3.16 0.20 0.68 - Trb 2.45 0.38 1.22 0.22 - Siv 5.65 1.52 1.24 1.37 1.53 - K-Br 2.63 1.29 2.38 1.29 0.74 3.75 - K- Mz 0.58 2.46 2.90 2.37 2.03 5.20 1.80 - Mil 3.81 9.35 12.24 8.98 8.28 14.70 8.59 5.07 - Bud 0.31 2.68 3.85 2.55 2.07 4.98 2.62 1.03 3.91 - Comparison of the mean values of measured traits (Diag. 1) shows a high degree of variation depending on the sampling point and significant discreteness. The greatest variation in traits is characteristic of the population from the village of Sivtsevo. A smaller length of elytra is typical of the ground beetles from points related to small- leaved forests: Gremyachevo, Tarusa, Bragina, and Ovrag Mozhayka in Kaluga, while a greater length is observed in beetles from broadleaf forests: Kireykovo, Trubetskoye, and Sivtsevo. Ground beetles from spruce forests show intermediate values for elytra length. The highest values of elytra length are observed in the village of Sivtsevo, and the lowest in Gremyachevo. The greatest width of elytra is found in the village of Sivtsevo, and the smallest in the village of Milyatino. The width of elytra is also somewhat greater in broadleaf forests. The greatest length of the pronotum is observed in the Kaluga urban forest, while the smallest is in Gremyachevo. Overall, the length of the pronotum shows higher values for ground beetles from spruce forests (Kaluga urban forest, Milyatino, Budnyansky), lower 728 Bull. Iraq nat. Hist. Mus. 18 (3): 721-738. Landscape-dependent variability values for deciduous forests, and average values for broadleaf forests. The greatest width of the pronotum is recorded in the village of Sivtsevo, and the smallest in Budnyansky. The length of the head of the ground beetles varies significantly; high values are observed in Milyatino, average values in Ovrag Mozhayka in Kaluga, Kireykovo, and Budnyansky, while other localities have uniformly low values. Distance between eyes has the highest values in Milyatino and the lowest in Sivtsevo. In general, ground beetles from spruce forests are characterized by higher values, while those from small-leaved forests have lower values; the values for ground beetles from broadleaf forests vary significantly. Ground beetles from certain localities show distinct differences. For example, ground beetles from the black alder swamp near Gremyachevo have low values for both elytra and pronotum lengths, while those from the linden forest near Sivtsevo are larger, show greater variation in trait sizes, and have low values for head length and distance between the eyes. Ground beetles from the spruce forest near Milyatino are characterized by a narrower body and greater values for head length and distance between eyes. Discriminant analysis of the beetles from different habitats of the locality “Bragino” “Bragino” locality is represented by three types of habitats with different vegetation types: willow-gray alder, black alder and linden. Analysis shows significant (F (12.11) = 23.13; p<0.001) and moderate discrimination (Wilks' Lambda: 0.64). The width of the elytra and pronotum does not make a significant contribution. Classification function coefficients show equally strong influence on discrimination by pronotum length and distance between the eyes. The greatest Mahalanobis distance (Tab. 3) is observed between the linden and black alder forests (small- and broadleaved), while the smallest distance is between the willow and black alder forests (both small-leaved). Table (3): Squared Mahalanobis distances (habitats in the locality Bragino), p<0.001. Willow Black alder Linden Willow - Black alder 0.89 - Linden 1.67 2.01 - 729 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Gorbunov et al. Diagram (1): Compared mean values of the measured traits of the beetle C. granulatus from different localities, boxes – mean error, whiskers – 1.96 mean error. The comparison of mean values (Diag. 2) shows that beetles from the linden forest are smaller in all traits except head length, while beetles from the willow and black alder forests have similar mean values for these traits. Beetles from the black alder stand exhibit greater head length, while the head length of beetles from the linden and willow forests is approximately equal. Diagram (2): Compared mean values of the measured traits of the beetle Carabus granulatus from different habitats of the locality Bragino; 1 – willow-gray alder, 2 – black alder, 3 – linden. Boxes – mean error, whiskers – 1.96 mean error. 730 Bull. Iraq nat. Hist. Mus. 18 (3): 721-738. Landscape-dependent variability Discriminant analysis by habitat type The results of the discriminant analysis by habitat type (Tab. 4) show weak (Wilks' Lambda= 0.81), but statistically significant (F (12.85) = 78.05; p < 0.001) differences among the beetles. The coefficients of the classification functions indicate that the parameters with the greatest significance for discrimination are the length of the pronotum, as well as the length and width of the elytra. Squared Mahalanobis distances are small but statistically significant (Tab. 3.). The maximum distance is observed between the morphometric parameters of ground beetles from spruce forest and small-leaved forest habitats (1.32), while the minimum distance (0.13) is found between beetles from the deciduous and broadleaf forests. Table (4): Squared Mahalanobis distances (habitat type), p<0.001 Broad-leaved forest Small-leaved forest Spruce forest Broad-leaved forest - Small-leaved forest 0.13 - Spruce forest 0.97 1.33 - Discriminant analysis by geomorphological units The results show significant and moderate differences between beetles from studied zones (Wilks' Lambda= 0.55, F (18.12) = 153.46, p<0.001). All traits make a statistically significant contribution to discrimination. According to coefficients of the classification functions, pronotum length has the highest discriminative value; other traits contribute to a lesser importance, while the contribution of pronotum width is very minimal. Squared Mahalanobis distances (Tab. 4.) indicate the greatest differences between the Moraine gently sloping plain of the Dnieper Glaciation with inherited post-Quaternary relief and the Flat glacial plain of the Moscow Glaciation. The smallest distance is observed between the Moraine gently sloping plain of the Dnieper Glaciation with inherited post-Quaternary relief and the Moraine gently undulating plain of the Moscow Glaciation. Table (4): Squared Mahalanobis distances (geomorphological units), p<0,001. I – Moraine- glacial gently undulating plain of the Dnieper Glaciation, II – Moraine gently sloping plain of the Dnieper Glaciation with inherited post-Quaternary relief, III – Moraine gently undulating plain of the Moscow Glaciation, IV – Flat glacial plain of the Moscow Glaciation. Zones I II III IV I - II 2.85 - III 1.50 1.06 - IV 1.39 5.17 3.73 - It can be seen (Diag. 3) that Kireykovo (region I, broadleaved forest) is closer to region IV (spruce forest) and region III (spruce and small-leaved forests), than to region II (broadleaved forest as well). Regions II and III are quite similar despite belonging to different vegetation 731 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Gorbunov et al. zones, but their geographic coordinates are close. Root 1 is mostly regulated by head length and distance between the eyes; while root 2 is shaped by length and width of the elytra, as well as pronotum length. Diagram (3): Ordination of morphometric features in different geomorphological units in the plane of two discriminant axes. Comparing Mahalanobis distances showed in Diag. (4) denotes the strongest diversification at the localities level; most of the distances fall within the range observed for the geomorphological units. Mahalonobis distances for the habitat type are the smallest, and in some cases, even smaller than the distances between spatially close populations within the Bragino locality. Diagram (4): Compared squared Mahalanobis distances between traits’ measures of the C. granulatus depending on the grouping variable for the discriminant analysis. 732 Bull. Iraq nat. Hist. Mus. 18 (3): 721-738. Landscape-dependent variability DISCUSSION The fact that discrimination of C. granulatus by measured traits is always reliable means that beetles have specific, significant differences in their morphology under various conditions. Although these differences are small and partly overlap with phenotypic variability, they are still consistent. Discrimination by habitat type based on dominant trees appeared to be very weak, while geographical units as a grouping variable showed much better discrimination. This likely means that the differences in populations of ground beetles are more significantly influenced by historical aspects of microevolutionary changes - caused by mechanisms such as genetic drift and the founder effect, followed by post-glacial changes - rather than by short-term phenotypic adaptation to different habitats. Dominant trees themselves have an indirect and limited effect on the beetle, as it is a generalist predator that requires prey, some shade, humidity, and cover in the form of grass or litter. Even on single locality level, discrimination based on dominant vegetation shows greater differences between populations than discrimination based on habitat type, which means that specific ecological features (microclimate, level of competition, position in landscape etc.) of each distinct ecosystem affect the morphometric traits more than average differences between habitat types. Coefficients of classification functions revealed that for both geomorphological and locality predictors, pronotum length has major significance, while pronotum width has very low contribution. This suggests that discrimination by locality type largely inherits the geomorphological patterns. Overall, at the provincial level, pronotum length has the greatest impact on discriminating beetles specimens. The shape of the pronotum is considered to be related to habitat usage and locomotion (Thiele, 1977); it influences forelimb mobility and the way beetles move through their environment (Talarico et al., 2007; Laparie et al., 2013; Jachertz et al., 2019). This is confirmed for different habitat types, where both pronotum length and width play relatively important roles in discrimination. Nevertheless, the low influence of pronotum width for locality and geomorphological grouping variables suggests that it reflects conservative, microevolutionary-based differences or functions as a proxy for body size, as elytra length does (Kaspari and Weiser, 1999). Overall, higher Wilks’ Lambda values for the discrimination by locality suggest that, at the locality level, various environmental and phylogenetic factors combine and provide more pronounced differences in morphological traits. Differences between populations from the locality Bragino are driven by distance between the eyes as well as pronotum length. It is known, that head width and the positioning of the eyes are correlated with predation behavior and lighting level (Bauer and Kredler, 1993; Bauer et al., 1998; Talarico et al., 2007). For the habitat-type level, beetles from spruce forests express higher distance between eyes than those from light small-leaved forests. 733 BULLETIN OF THE IRAQ NATURAL HISTORY MUSEUM Gorbunov et al. In terms of the morphometric structure of populations, the central position is occupied by the population from Kireykovo, located in a minimally disturbed area of broadleaf forests on a watershed, which can be considered a reference for the broadleaf forest zone. The population from Budnyansky, which belongs to the coniferous-broadleaf forest zone and is situated within a raised bog, is the closest to it. Formally, it is represented by a spruce forest, which, however, is very rich in broadleaved forest inhabitants (Aleksanov et al., 2024a), characteristic of morainic islands. It is interesting to note that despite the reduced abundance of beetles (Aleksanov et al., 2024b); a balanced morphometric structure has been preserved here. Another group is formed by populations in the eastern part of the Kaluga region (broadleaf forest zone), associated with the Oka River valley (Bragino, Gremyachevo, Tarusa), to which a watershed locality (Trubetskoye) is added. Beetles in these localities are smaller in most traits (or lengths). On one hand, this can be explained by the poverty of infertility sediments with a high sand content, which are poor both in terms of moisture and mineral nutrients (compared to morainic deposits). The lower favorability of such habitats for beetles may result from poorer shelter quality (less deadwood for wintering, less dense herbaceous cover) as well as a poorer food base (fewer mollusks, earthworms). On the other hand, a no less important factor could be the prolonged economic use of the forests (agricultural, park, forestry, recreational), leading to their fragmentation and degradation of the soil cover. The negative reaction of the species to recreational load has been demonstrated for this region (Sionova, Alekseev, 2024). A special case is the park in the village of Sivtsevo, where the beetles are on average the largest, which can be explained by the favorable herbaceous cover, but the variability is very high. Overall, all fragmented forests in the eastern part of the Kaluga region differ significantly from each other in individual morphometric traits. This may be related to selection for specific habitat conditions, but it is more likely a consequence of the small number of individuals that colonized these habitats (likely resulting in reduced genetic diversity). The forests of Kaluga occupy an intermediate position between wild, quasi-climax forests and economically disturbed forests in terms of morphometric structure. They are smaller, but maintain the morphological proportions of beetles from quasi-climax forests. Despite being within the boundaries of a present-day city, they have remained forests for a long time, with moderate anthropogenic impact. Compared to broadleaf forests, they are less fertile—both climatically and in terms of soil-forming rocks. At the opposite end of the virtual morphometric space is the wet spruce forest in the western part of the Kaluga region (Milyatino), it is wet, with moderate disturbance (apparently, it began to overgrow after World War II), but the glacial sediments from the Moscow glaciation are less fertile than those in Kireykovo. 734 Bull. Iraq nat. Hist. Mus. 18 (3): 721-738. Landscape-dependent variability At the locality level, the linden forest in the ravine is harsher because it is a dry, young forest with hairy sedge, where beetles show reasonably smaller morphometric traits. CONCLUSIONS The study showed that C. granulatus beetles classified by various grouping factors have significant but relatively weak differences in their morphological traits. Pronotum length is a key trait for distinguishing populations, while pronotum width has low significance. Morphological differences in populations of C. granulatus are primarily influenced by historical microevolutionary processes, such as genetic drift, rather than being solely the result of phenotypic adaptation to habitats. Differences between beetles from various ecosystems depend on local ecological factors, such as microclimate, rather than just habitat type. At both spatial levels, similarity in morphometric structure does not directly correlate with geographic distance. between points on the ground. Populations from the eastern part of the Kaluga region are smaller, which may be related to poor soil quality and intensive use of forests. Fragmented forests exhibit significant differences in morphometric traits, likely due to the small number of colonizing individuals. Overall, the results highlight the importance of considering phylogenetic and ecological factors when studying morphological differences. AKNOWLEDGEMENTS We would like to thank S.K. Alekseev, M.P. Shashkov, M.I. Garkunov, D.V. Khvaletsky, S.E. Carpukhin and V.V. Perov for the help with sampling carabid beetles. CONFLICT OF INTEREST STATEMENT This study is funded by the State Task Theme 730000P.16.1.OH17AA106000 “Biological diversity of Eastern Europe under the influence of natural and climatic factors in historical and modern contexts”. LITEREATURE CITED Aleksanov, V. V., Alekseev, S. K., Ruchin, A. B. and Esin, M. N. 2024 a. Carabid beetles under the influence of megafires in pine and secondary forests of Central European Russia. Diversity, 16(7): 370. [CrossRef] Aleksanov, V. V., Alekseev, S. K., Garkunov, M. I., Karpukhin, S. E., Prokhorova, N. E., Perov, V. V., Teleganova, V. V., Khvaletsky, D. V. and Shmytov, A. A. 2024 b. 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Carabus granulatus L., 1758 للنوع البيئية االنماط الرتبط التغايرالورفومتري (Coleoptera, Carabidae) بيردنيك*، سيرجي ريفغاتشاجيدولين*، ألكسانوف**، فيكتور غوربونوف*، رومان سوخودولسكايا* ورايسا و التحاد للعلوم، تتارستان لكاديمية التابع العدنية الثروة واستخدام البيئة مشاكل *معهد قازان. ،420087 اللووسي، بوريسوفكا. قلية ،309342 اللووسي، التحاد الحكومية، الطبيعية بيلوجوريي **محمية ،248000 اللووسي، التحاد الحدائق"، "مديلية كالوغا لنطقة الحكومية اليزانية ***مؤسسة كالوغا. قازان. ،420012 اللووسي، التحاد الحكومية، الطبية قازان ****جامعة قلية ،422623 اللووسي، التحاد الطبيعي، الحيوي للمحيط الحكومية فولغا-كاما *****محمية سادوفي. 2025/6/20 النشر: ،2025/5/29 القبول: ،2025/5/24 2025/2/28،الراجعة: استتلم: اخالصة جماعـــــــــــات فــــي الشكلي التغايل مظاهل استقصاء إلى الدراســــــــــــــــــــــة هــــــــــــــذه هدفت ضمن النتشلة Carabidae عائلة و Coleoptera رتبة من Carabus granulatus L., 1758 على الورفولوجي التكيف ديناميات فهم إطار في وذلك كالوغا، إقليم في متباينة بيئات فلوق عن الكشف إلى التغيرات متعدد التمييزي التحليل أففسى وقد الحلي. الستوى أن ن تبيي حيث الدروسة، الجماعات بين دقيقة، كانت وإن إحصائية، دللة ذات شكلية بينها. ا تمييزز الكثر السمة عد يي )البرونوتوم( المامي الصدر طول التكيف إلى الول القام في عزى يي ل الورفومتري التغايل أن النتائج أظهلت لقد التي الدقيقة التطورية بالعمليات بعيد حدي إلى يلتبط بل الوئل، نوع مع الباشل الفينوتيبي البيئية العوامل أن العطيات نت بيي كما الحلية. للسالت اليكلوتطوري التاريخ عبر نشأت 738 Bull. Iraq nat. Hist. Mus. 18 (3): 721-738. Landscape-dependent variability أكثر ا تأثيرز تمتلك الوارد، وفلة وتفاوت الحلي الناخ في الدقيقة كالتغيرات الوضعية، عام. نحوو على بتصنيفالوائل ز مقارنة الشكلية السمات تشكيل في فاعلية القليم شلق في الواقعة الجماعات سمت ي ات فقد الجغلافي، التوزيع حيث من أما الحلاجي الستغال عن الناتج الضغط إلى يعود ذلك أن ح ّي ويل الكلي، الحجم بانخفاضفي ا تمايزز أظهلت الجزأة الغابات في القاطنة الجماعات أن الدراسة كشفت كما ف. ي الكّ جماعات تأسيس عن الناتج الؤسس” “تأثير عن ناجم أنه حتمل يي ا، واضحز ا مورفومتريز الفلاد. من محدود عدد من جديدة