Morphometric diversity of Zabrus morio (Coleoptera: Carabidae) in relation to sex and altitude in Uzbekistan's mountain ecosystems

Main Article Content

DILNOZA F. ZOKIROVA
JASUR A. KUDRATOV
RAFIK A. KHAMZAEV
ASOLAT B. KHAMIDOVA
BEKSOD OTAKULOV
GULRUH URINOVA
IRODA ZOHIDOVA
ERKIN ABDULLAEV
FARIZA RAYIMOVA
ABDUVAET PAZILOV
SHAHLO TURSUNOVA
MUNISA BAKHRILLAYEVA
FAZLITDIN Z. KHALIMOV

Abstract

Abstract. Zokirova DF, Kudratov JA, Khamzaev RA, Khamidova AB, Otakulov B, Urinova G, Zohidova I, Abdllaev E, Rayimova F, Pazilov A, Tursunova S, Bakhrillayeva M, Khalimov FZ. 2025. Morphometric diversity of Zabrus morio (Coleoptera: Carabidae) in relation to sex and altitude in Uzbekistan's mountain ecosystems. Biodiversitas 26: 4730-4745. Zabrus morio is the only representative of the genus Zabrus in Uzbekistan, considered a potential pest and widely distributed in mountainous and foothill ecosystems. Studying its morphometric characteristics is important for understanding population traits and adaptive responses. A total of 115 individuals (50 females, 65 males) from 10 sites along an altitudinal gradient were examined. Sites above 1,200 masl were represented by mountain steppes, subalpine meadows, juniper woodlands, and alpine meadows. Sites below 1,200 m were characterized by arid ephemeral-grass steppes, sagebrush-ephemeral steppes, and shrub-steppe communities. Ten morphometric traits were measured using a binocular microscope. Head length was the most variable trait (CV = 10.2%), while elytral width was the least variable (CV = 5.5%); in general, body length parameters were more variable than widths. The strongest correlation was found between elytra length and total body length (r = 0.93), while head length and pronotum length were the most independent traits (r = 0.49). Females showed greater variability in head dimensions and pronotum length, while males exhibited higher variability in elytral width and interocular distance. In females, elytra length correlated most strongly with total body length (r = 0.92), whereas in males, head width closely correlated with pronotum width (r = 0.95), which may indicate a relationship between head size and pronotum size. Overall, trait variability decreased with increasing altitude, with only male head length showing a statistically significant reduction. These findings emphasize the adaptive morphometric variability of Z. morio depending on sex and ecological conditions, as well as the potential implications of these correlations for the species’ ecology and evolution.

Article Details

Section

Articles

References

Abdurakhmanov GM, Kassem AS. 2013. The role and place in agroecosystems of soil beetles (Coleoptera: Carabidae, Scarabaeid?e, Elateridae, Tenebrionidae) - pests of agricultural crops in the Republic of Dagestan. Sci J Kuban State Agrarian Univ 86: 1-11. [Russian]

Abdurakhmanov GM, Klicheva SM. 2010. Zoogeographical characteristics of the ground beetles (Coleoptera, Carabidae) of the Steppe areas of the Southeast of Russia and Northeast of Azerbaijan. South Russia: Ecol Dev 1: 63-75. [Russian]

Alcantara MJM, Fontanilla AM, Ashton LA, Burwell CJ. 2024. Bugs and Bergmann's rule: A cross-taxon large-scale study reveals idiosyncratic altitudinal and latitudinal body size patterns for different insect taxa. Entomol Gen 44 (3): 715-725. DOI: 10.1127/entomologia/2024/2246.

Ananina TL, Sukhodolskaya RA. 2019. Study of the morphometric structure of ground beetle populations of the Barguzin Ridge on the example of Carabus odoratus barguzinicus (Shil, 1996). In: Natural complexes of the North-Eastern Transbaikalia. BNTs SB RAS Publishers, Russia. [Russian]

Anderegg LDL. 2023. Why can't we predict traits from the environment? New Phytol 237: 1998-2004. DOI: 10.1111/nph.18586.

Baranovská E, Knapp M. 2018. Steep converse Bergmann's cline in a carrion beetle: Between- and within-population variation in body size along an elevational gradient. J Zool 304: 243-251. DOI: 10.1111/jzo.12527.

Baranovská E, Tajovský K, Knapp M. 2019. Changes in the body size of carabid beetles along elevational gradients: A multispecies study of between- and within-population variation. Environ Entomol 48 (3): 583-591. DOI: 10.1093/ee/nvz036.

Budilov PV. 2013. Sexual dimorphism of body size in a ground beetle species (Notiophilus reitteri Spaeth). Reg Probl 16 (1): 49-54. [Russian]

Bui VB, Ziegler Th, Bonkowski M. 2020. Morphological traits reflect dung beetle response to land use changes in tropical karst ecosystems of Vietnam. Ecol Indic 108: 105697. DOI: 10.1016/j.ecolind.2019.105697.

Bulgarella M, Trewick SA, Godfrey AJ, Sinclair BJ, Morgan-Richards M. 2015. Elevational variation in adult body size and growth rate, but not in metabolic rate, in the tree weta Hemideina crassidens. J Insect Physiol 75: 30-38. DOI: 10.1016/j.jinsphys.2015.02.012.

Desender K. 1986. Distribution and ecology of carabid beetles in Belgium (Coleoptera: Carabidae). Part 2. Rijksuniversiteit Gent.

Dudarova HYu, Abdurakhmanov ShG. 2009. Distribution of beetle pests among agricultural crops, their trophic links with the vegetation of natural landscapes. South Russia: Ecol Dev 2: 108-115. [Russian]

Favé MJ, Johnson RA, Cover S, Handschuh S, Metscher BD, Müller GB, Gopalan S, Abouheif E. 2015. Past climate change on Sky Islands drives novelty in a core developmental gene network and its phenotype. BMC Evol Biol 15: 1-21. DOI: 10.1186/s12862-015-0448-4.

Fountain-Jones NM, Baker SC, Jordan GJ. 2015. Moving beyond the guild concept: Developing a practical functional trait framework for terrestrial beetles. Ecol Entomol 40: 1-13. DOI: 10.1111/een.12158.

Hodkinson ID. 2005. Terrestrial insects along elevation gradients: Species and community responses to altitude. Biol Rev 80 (3): 489-513. DOI: 10.1017/S1464793105006767.

Ji Q, Xie Zh, Wu Y, Wan Zh, Xu C, Wu D, Chen T-W, Ordonez A. 2024. Intraspecific trait variation of carrion beetle species and communities across elevations. Insect Conserv Divers 17: 1113-1126. DOI: 10.1111/icad.12772.

Karpova VE. 1992. Efficiency of catching ground beetles (Coleoptera, Carabidae) with different types of Barber traps. Biol Sci 5: 84-88.

Kaspari M, Marshall KE, Weiser MD, Siler CD, Theriot MK, de Beurs K. 2024. Geographic gradients in a functional trait: Drivers of body size and size diversity of ground invertebrate communities. Ecosphere 15: Article e4785. DOI: 10.1002/ecs2.4785.

Kawano K. 2016. Comparative quantification of intralocational, interlocational, and interspecific variability in stag beetles (Coleoptera: Lucanidae) and the questions of phenotypic plasticity and species selection. Ann Entomol Soc Am 109 (4): 555-566. DOI: 10.1093/aesa/saw011.

Khalimov F, Rakhimov M, Khamzaev R, Abdullaev E, Usanov U. 2023. Composition and structure of the entomofauna of Ferula (Ferula kuhistanica) in different sections of the Zarafshan Ridge. J Entomol Res Soc 25 (2): 275-286. DOI: 10.51963/jers.2023.86.

Khalimov F. 2020. The ground beetles (Coleoptera, Carabidae) of the Karatepa and Chakilkalyan mountains (west part of Zarafshan Mountains Range, Uzbekistan). Biosyst Divers 28 (3): 265-271. DOI: 10.15421/012035.

Khalimov F. 2023. Composition and structure of the fauna of ground beetles (Coleoptera, Carabidae) of the Zerafshan Range. Acta Biol Sibirica 9: 113-125. DOI: 10.5281/zenodo.7725474.

Khojakulov D, Khaydarov Kh, Rabbimov A, Mukimov T, Matkarimova G, Davronkulova F, Ochilov U, Alikulov BS. 2024. Biological, physiological and economic characteristics of Onobrychis chorasanica Bunge ex Bois. (Sainfoin) under sowing conditions. Plant Sci Today 11 (3): 14-21. DOI: 10.14719/pst.3180.

Koçak Y, Do?an Sarikaya A, Sarikaya Ö, Macirella R, Talarico F, Brunelli E. 2025. Head morphology in three species of tiger beetles (Coleoptera: Cicindelidae): A geometric morphometric study. Eur Zool J 92 (1): 496-506. DOI: 10.1080/24750263.2025.2485102.

Koivula SM. 2011. Evolution of insect life histories in relation to time constraints in seasonal environments: Polymorphism and clinal variation. Acta University of Oulu 569: 1-70.

Komlyk V, Brygadyrenko V. 2020. Morphological variability of Bembidion varium (Coleoptera, Carabidae) in a gradient of soil salinity. Folia Oecol 47 (1): 23-33. DOI: 10.2478/foecol-2020-0004.

Komlyk VO, Brygadyrenko VV. 2019. Morphological variability of Bembidion aspericolle (Coleoptera, Carabidae) populations in conditions of anthropogenic impact. Biosyst Divers 27 (1): 21-25. DOI: 10.15421/011903.

Kotze DJ, Brandmayr P, Casale A, Dauffy-Richard E, Dekoninck W, Koivula MJ, Lövei GL, Mossakowski D, Noordijk J, Paarmann W, Pizzolotto R. 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. DOI: 10.3897/zookeys.100.1523.

Kudratov J, Pazilov A, Zokirova D, Bekzod Otakulov B, Jalilov J, Urinova X, Khalimov F, Izzatullaev Z. 2024. Complexes of terrestrial molluscs of various biotopes in the Gissar Range, Uzbekistan. Acta Biol Sibirica 10: 349-359. DOI: 10.5281/zenodo.11160044.

Li XD, Jiang GF, Li R, Bai Y, Zhang GS, Xu SJ, Deng WA. 2024. Molecular strategies of the pygmy grasshopper Eucriotettix oculatus adapting to long-term heavy metal pollution. Ecotoxicol Environ 276: 116301. DOI: 10.1016/j.ecoenv.2024.116301.

Liu S, Tong J, Xu M, Meng Q, Shi Y, Zhao H, Li Y. 2024. The effect of elevation gradient on distribution and body size of carabid beetles in the Changbaishan Nature Reserve in Northeast Asia. Insects 15: 688. DOI: 10.3390/insects15090688.

Löbl I, Löbl D. 2017. Catalogue of Palaearctic Coleoptera, 1: Archostemata - Myxophaga - Adephaga. Revised and Updated Edition, BRILL, Leiden, Boston.

Lorenz C, Almeida F, Almeida-Lopes FL, Louise C, Pereira SN, Petersen V, Vidal PO, Virginio F, Suesdek L. 2017. Geometric morphometrics in mosquitoes: What has been measured? Infect Genet Evol 54: 205-215. DOI: 10.1016/j.meegid.2017.06.029.

Lövei GL, Sunderland KD. 1996. Ecology and behavior of ground beetles (Coleoptera: Carabidae). Annu Rev Entomol 41: 231-256. DOI: 10.1146/annurev.en.41.010196.001311.

Luzyanin S, Saveliev A, Ukhova N, Vorobyova I, Solodovnikov I, Anciferov A, Shagidullin R, Teofilova T, Nogovitsyna S, Brygadyrenko V, Alexanov V, Sukhodolskaya RA. 2022b. Modeling sexual differences of body size variation in ground beetles in geographical gradients: a case study of Pterostichus melanarius (Illiger, 1798) (Coleoptera, Carabidae). Life 12: 112. DOI: 10.3390/life12010112.

Luzyanin SL, Gordienko TA, Saveliev AA, Ukhova NL, Vorobeva IG, Solodovnikov IA, Anciferov AA, Nogovitsyna SN, Aleksanov VV, Teofilova TM, Sukhodolskaya RA. 2022a. Impact of climatic factors on sexual size dimorphism in ground beetle Pterostichus melanarius (Illiger, 1798) (Coleoptera, Carabidae). Ecol Montenegrina 58: 1-13. DOI: 10.37828/em.2022.58.1.

Magura T, Lövei GL. 2020. The permeability of natural versus anthropogenic forest edges modulates the abundance of ground beetles of different dispersal power and habitat affinity. Diversity 12 (9): 320. DOI: 10.3390/d12090320.

Magura T, Lövei GL. 2021. Consequences of urban living: Urbanization and ground beetles. Curr Landsc Ecol Rep 6 (1): 9-21.

Mamanov SS, Rakhimov MR, Khalimov FZ, Khamzaev RA, Abdullayev EN, Jabborov AR, Khamidova AB, Shodmonov FA, Ashrapov AA, Ochilbekova S. 2024. Review of the genus Dioctria Meigen, 1803 (Diptera: Asilidae) in Uzbekistan with the description of two new species. Acta Biol Sibirica 10: 1543-1557. DOI: 10.5281/zenodo.14303122.

Marshall J, Miller M, Lelito J, Storer A. 2013. Latitudinal variation in body size of Agrilus planipennis and relationship with fecundity. Agric For Entomol 15: 294-300. DOI: 10.1111/afe.12017.

Medetov MZH, Abdullaeva JK, Reymov A, Miratdinova AM, Seytmuratov AK, Tajibaeva JD, Kadirov RS, Utemuratov NA, Kimyonazarov SK, Kudratov J, Urazova RS, Keldiyorova XX, Uralov UB. 2025. Diversity of true bugs (Hemiptera: Heteroptera) of the Southern Aral Sea Region, Uzbekistan. Biodiversitas 26 (7): 3125-3135. DOI: 10.13057/biodiv/d260704.

Messier J, McGill BJ, Enquist BJ, Lechowicz MJ. 2017. Trait variation and integration across scales: Is the leaf economic spectrum present at local scales? Ecography 40: 685-697. DOI: 10.1111/ecog.02006.

Montejo-Kovacevich G, Salazar PA, Smith SH, Gavilanes K, Bacquet CN, Chan YF, Jiggins CD, Meier JI, Nadeau NJ. 2021. Genomics of altitude-associated wing shape in two tropical butterflies. Mol Ecol 30: 6387-6402. DOI: 10.1111/mec.16067.

Moran EV, Hartig F, Bell DM. 2016. Intraspecific trait variation across scales: Implications for understanding global change responses. Global Change Biol 22: 137-150. DOI: 10.1111/gcb.13000.

Mukhtorova S, Alikulov B, Yuldosheva M, Maxammadieva D, Khidirova U, Kabulova F, Ismailov Z. 2024. Diversity of endophytic bacteria isolated from Peganum harmala distributed in arid regions in Uzbekistan. Regulatory Mechanisms Biosyst 15 (2): 286-291. DOI: 10.15421/022441.

Muscarella R, Uriarte M. 2016. Do community-weighted mean functional traits reflect optimal strategies? Proc Royal Soc B 283: 20152434. DOI: 10.1098/rspb. 2015.2434.

Naydenov AE, Makhov IA, Mironov VG, Rakhimov MR, Khalimov FZ, Yakovlev RV. 2025. Materials on the Lepidoptera of Uzbekistan. Part I. Family Geometridae Leach, 1815. Acta Biol Sibirica 11: 589-611. DOI: 10.5281/zenodo.15274938.

Nishikawa M, Ikeda H, Kubota K, Sota T. 2010. Taxonomic redefinition and natural history of the endemic silphid beetle Silpha longicornis (Coleoptera: Silphidae) of Japan, with an analysis of its geographic variation. Zootaxa 2648: 1-31. DOI: 10.11646/zootaxa.2648.1.1.

Ornaghi S, Valle B, Caccianiga M, Seppi R, Gobbi M. 2023. Sex-ratio and body size plasticity in two cold-adapted ground beetles co-occurring in a periglacial area of the European Alps (Coleoptera: Carabidae). Fragm Entomol 55 (1): 45-56. DOI: 10.13133/2284-4880/1442.

Parhomenko O, Langraf V, Petrovi?ová K, Komlyk V, Brygadyrenko V. 2022. Morphometric variability of ground beetles Bembidion minimum (Coleoptera, Carabidae): who should change more, males or females? Nat Conserv Res 7 (1): 42-69. DOI: 10.24189/ncr.2022.008.

Polat A. 2023. A study on morphological variations of male Helophorus (Helophorus) aquaticus (L., 1758) (Coleoptera: Helophoridae) in Türkiye. Turkish J Entomol 47: 59-72. DOI: DOI: 10.16970/entoted.1192141.

Rakhimov MR. 2023. Checklist of hover flies (Diptera, Syrphidae) of the western part of the Zarafshan mountain ridge. Acta Biol Sibirica 9: 167-193. DOI: 10.5281/zenodo.7835401.

Rusynov VI, Brygadyrenko VV. 2017. Morphological variability of a population of Anatolica eremita (Coleoptera, Tenebrionidae): constancy of morphometric indices with variation of linear parameters of the body. Baltic J Coleopterol 17 (2): 205-217.

Sazhnev AS, Ermakov NM, Kondratyev EN. 2024. Beetles (Coleoptera) from the burrows of the red-tailed gerbil Meriones libycus Lichtenstein, 1823 (Mammalia: Muridae) in the south-western Turkmenistan. Ecosyst Transformation 7 (2): 208-222. DOI: 10.23859/estr-230316.

Shelomi M. 2012. Where are we now? Bergmann's Rule sensu lato in insects. Am Nat 180 (4): 511-519. DOI: 10.1086/667595.

Shelomi M. 2023. A practical guide to collections-based research on ecogeography. Ecol Evol 13 (1): e10211. DOI: 10.1002/ece3.10211.

Smith JML, Catherall-Ostler AM, Mashoodh R, Kilner RM. 2024. Sexual dimorphism in head size in wild burying beetles. Ecol Entomol 49: 846-856. DOI: 10.1111/een.13359.

Stachewicz JD, Fountain-Jones NM, Koontz A, Woolf H, Pearse WD, Gallinat AS. 2021. Strong trait correlation and phylogenetic signal in North American ground beetle (Carabidae) morphology. Ecosphere 12 (11): e03832. DOI: 10.1002/ecs2.3832.

Stanbrook RA, Harris WE, Wheater CP, Jones M. 2021. Evidence of phenotypic plasticity along an altitudinal gradient in the dung beetle Onthophagus proteus. PeerJ 9: e10798. DOI: 10.7717/peerj.10798.

Stillwell RC. 2010. Are latitudinal clines in body size adaptive? Oikos 119: 1387-1390. DOI: 10.1111/j.1600-0706.2010.18670.x.

Sugahara R, Tanaka S. 2018. Environmental and hormonal control of body color polyphenism in late-instar desert locust nymphs: Role of the yellow protein. Insect Biochem Mol Biol 93: 27-36. DOI: 10.1016/j.ibmb.2017.12.004.

Sukhodolskaya R. 2013. Intraspecific body size variation in ground beetles (Coleoptera, Carabidae) in urban - suburban - rural - natural gradient. Acta Biologica Universitatis Daugavpiliensis 13 (1): 121-128.

Sukhodolskaya RA, Ananina TL. 2015. Altitudinal variation in population density, body size, and morphometric structure in Carabus odoratus Shil, 1996 (Coleoptera: Carabidae). Acta Biol Universitatis Daugavpiliensis 15: 179-190.

Sukhodolskaya RA, Saveliev AA. 2014. Effects of ecological factors on size-related traits in the ground beetle Carabus granulatus L. (Coleoptera, Carabidae). Russian J Ecol 45 (5): 414-420. [Russian]

Sukhodolskaya RA, Saveliev AA. 2017. Geographi?al variation of sexual size dimorphism of the ground beetle Carabus granulatus L. (Coleoptera, Carabidae). Ecol Nat Syst 4: 3-10. [Russian]

Sun X, Xu M, Jia L. 2023. Responses of diversity and morphometric traits of ground beetles to soil chemical properties in an industrial city of China. Ecol Indic 154: 110575. DOI: 10.1016/j.ecolind.2023.110575.

Talarico F, Mazzei A, Gangale C, Scrivano G, Brandmayr P. 2020. Morphometric differences in populations of Nebria kratteri Dejean & Boisduval, 1830 from two old forests in Calabria (Coleoptera, Carabidae). Fragm Entomol 52 (1): 57-62. DOI: 10.13133/2284-4880/408.

Venn S. 2007. Morphological responses to disturbance in wing-polymorphic carabid species (Coleoptera: Carabidae) of managed urban grasslands. Baltic J Coleopterol 7 (1): 51-60.

Wos G, Palomar G, Marsza?ek M, Babik W, Sniegula S. 2023. The effect of temperature and invasive alien predator on genetic and phenotypic variation in the damselfly Ischnura elegans: Cross-latitude comparison. Front Zool 20: 13. DOI: 10.1186/s12983-023-00494-z.

Yadav S, Stow AJ, Dudaniec RY. 2019. Detection of environmental and morphological adaptation despite high landscape genetic connectivity in a pest grasshopper (Phaulacridium vittatum). Mol Ecol 28: 3395-3412. DOI: 10.1111/mec.15146.

Yang J, Lu J, Chen Y, Yan E, Hu J, Wang X, Shen G. 2020. Large underestimation of intraspecific trait variation and its improvements. Front Plant Sci 11: 53. DOI: 10. 3389/fpls.2020.00053.

Zhang Y, Xu H, Wang Z, Jie H, Gao F, Cai M, Wang K, Chen D, Guo R, Lin Z, Niu Q. 2023. A key gene for the climatic adaptation of Apis cerana populations in China, according to selective sweep analysis. BMC Genomics 24: 100. DOI: 10.1186/s12864-023-09167-x.

Zokirova DF, Alimova LH, Khalimov FZ. 2022. Study of morphometric features of Machozetus lehmanni Menetries, 1848-endemic of Central Asia. Sci Rev Biol Sci 1: 16-21. DOI: 10.17513/srbs.1253. [Russian]

Zokirova DF, Khalimov FZ. 2022. Morphometric features of the beetle Acinopus (Acinopus) laevigatus Menetries, 1832 (Coleoptera, Carabidae) in the mountain ecosystems of Uzbekistan. Bull Iraq Nat Hist Museum 17 (2): 141-153. DOI: 10.26842/binhm.7.2022.17.2.0141.

Most read articles by the same author(s)