Article
| Article name | PROGLACIAL SUCCESSIONS OF SPRINGTAIL ASSEMBLAGES (COLLEMBOLA) ALONG RETREATING GLACIERS IN KABARDINO-BALKARIA, GREATER CAUCASUS, RUSSIA |
| Authors | Maria D. Antipova, Junior Researcher, Laboratory of Synecology, Severtsov Institute of Ecology and Evolution, Russian Academy of Sciences, (119071, Russia, Moscow, Leninsky Prospekt, 33); iD ORCID: https://orcid.org/0000-0002-3517-9720; e-mail: antimar.2410@gmail.com |
| Reference to article | Antipova M.D., Bushueva I.S., Babenko A.B. 2026. Proglacial successions of springtail assemblages (Collembola) along retreating glaciers in Kabardino-Balkaria, Greater Caucasus, Russia. Nature Conservation Research 11(1): 71–92. https://dx.doi.org/10.24189/ncr.2026.006 Electronic Supplement 1. Additional data for the paper by Antipova et al. (2026) (Link) |
| Section | Research articles |
| DOI | https://dx.doi.org/10.24189/ncr.2026.006 |
| Abstract | Since the end of the Little Ice Age, glacier retreat has been recorded globally, with its rates steadily increasing. Glacier forelands serve as convenient areas for studying the patterns of biotic community formation during primary succession. Collembola (hereinafter – springtails) typically play key roles in primary successions, being among the first colonists of territories newly freed from ice. The present study focuses on successional changes in springtail assemblages in the valleys of the Cherek-Khulamsky (Bezengi) and Adylsu along the Bezengi and Kashkatash glacier forelands, respectively (Kabardino-Balkaria, northern Caucasus). The Cherek-Khulamsky valley is located within the Kabardino-Balkarian High-Mountain State Nature Reserve, while the Adylsu valley is part of the Prielbrusye National Park. The objective of the present study is characterising the species composition and structure of springtail assemblages along the glacier chronosequences, as well as identifying the local features of the dynamics of these communities. Using standard soil zoological methods, we examined springtail communities across ten precisely dated plots (1–150 years since deglaciation) in both study valleys. On both locations, pioneer springtail complexes formed immediately after glacier retreat include specialised species, such as representatives of the nivalis group of the genus Desoria, which are replaced by widely distributed forms within 3–5 years. The dynamics of successional processes vary significantly between the Bezengi and Kashkatash glacier forelands. At Bezengi, the species richness has gradually increased, reaching a maximum after 135 years, whereas at Kashkatash, a sharp diversity increase is observed as early as 74 years since glacier retreat, at the shrub overgrowth stage. Springtail density also varies: at Bezengi it fluctuates from 36 individuals/dm2 to 177 individuals/dm2, reaching the maximum values just 14 years after deglaciation (grassland stage), while at Kashkatash the first noticeable population increase up to 632 individuals/dm2 is recorded on 74-year old surfaces, with an absolute maximum (726 individuals/dm2) achieved in 150-year old pine forests. The Kashkatash glacier foreland shows more pronounced differentiation between early (pioneer) and late (forest) successional stages than that of Bezengi, evident in both population abundance and species composition. At the same time, radical changes in springtail communities are observed in both valleys during transitions either from grasslands to shrublands or from shrublands to forests. Comparison with similar studies reveals a great role of the local features of a particular foreland. Regional differences could be accounted for by microclimatic, geomorphological and biotic factors. At Bezengi, where subalpine grasslands subject to anthropogenic influence predominate, succession seems to proceed more smoothly, while at Kashkatash the development of pine forests creates stable conditions favouring the formation of sustainable communities. Orographic features, such as slope steepness and the presence of rigels, also render a substantial impact, determining the environmental patchiness and the rate of successional changes. The data obtained contribute to understanding the patterns of biotic community formation in montane ecosystems under deglaciation and emphasise the need for further comparative studies to predict changes under global warming conditions. |
| Keywords | arthropods, chronosequence, climate change, glacier forelands, montane ecosystem, pioneer species, primary succession |
| Artice information | Received: 28.08.2025. Revised: 18.11.2025. Accepted: 06.01.2026. |
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Antipova M.D., Babenko A.B. 2022. An annotated checklist of the springtails (Hexapoda: Collembola) recorded from the foothill and mountain parts of the Republic of North Ossetia–Alania, North Caucasus, Russia. Russian Entomological Journal 31(4): 331–345. DOI: 10.15298/rusentj.31.4.01 Antipova M.D., Babenko A.B. 2023. Methods of collecting springtails in conditions of mountain glacier melting. In: Environmental safety and conservation of plant and animal genetic resources in Russia and adjacent territories. Vladikavkaz: North-Ossetian State University. P. 15–22. [In Russian] Antipova M.D., Babenko A.B. 2024. The formation of springtail assemblages (Hexapoda, Collembola) along the retreating Tsey Glacier, North Ossetia-Alania. Biology Bulletin 51(8): 2401–2419. DOI: 10.1134/S1062359024701139 Babenko A.B. 1985. Comparison of different sampling methods for atmobiont Collembola. In: M.S. Gilyarov (Ed.): Problems of soil zoology. Ashkhabat: Institute of Zoology of the Turkmen SSR Academy of Sciences. P. 22–24. [In Russian] Babenko A.B., Ponomarev A.V. 2023. Spiders (Aranei) of the periglacial landscapes of the Tseiskii Gorge, North Ossetia-Alania, Caucasus, Russia. Biology Bulletin 50(9): 2172–2185. DOI: 10.1134/S1062359023090054 Bondarenko S.V. 2011. The vegetation of the gorge Bezengy (the Kabardino-Balkar State Nature Reserve, the Central Caucasus). In: Problems of botany of Southern Siberia and Mongolia. Barnaul: AltSU Publishing House. P. 16–18. [In Russian] Buda J., Azzoni R.S., Ambrosini R., Franzetti A., Zawierucha K. 2020. Effects of locality and stone surface structure on the distribution of Collembola inhabiting a novel habitat – the stone-ice border on an alpine glacier. Acta Oecologica 108: 103629. DOI: 10.1016/j.actao.2020.103629 Bushueva I.S. 2013. Glacier fluctuations in the Central and Western Caucasus over the last 200 years based on cartographic, historical and bioindication data. PhD Thesis. Moscow: Institute of Geography RAS. 169 p. [In Russian] Bushueva I.S., Solomina O.N. 2012. Kashkatash Glacier fluctuations in the XVII–XXI centuries from cartographic, dendrochronological and lichenometric data. Ice and Snow 52(2): 121–130. DOI: 10.15356/2076-6734-2012-2-121-130 [In Russian] Chernov A.V., Kuznetsova N.A., Potapov M.B. 2010. Springtail Communities (Collembola) of Eastern European Broad-leaf Forests. Entomological Review 90(5): 556–570. DOI: 10.1134/S0013873810050039 Cody M.L. 1975. Towards a theory of continental species diversities: bird distributions over Mediterranean habitat gradients. In: M.L. Cody, J.M. Diamond (Eds.): Ecology and Evolution of Communities. Cambridge: Harvard University Press. P. 214–257. Crosta A., Valle B., Caccianiga M., Gobbi M., Ficetola F.G., Pittino F., Franzetti A., Azzoni R.S., Lencioni V., Senese A., Corlatti L., Buda J., Poniecka E., Novotná Jaroměřská T., Zawierucha K., Ambrosini R. 2025. Ecological interactions in glacier environments: a review of studies on a model Alpine glacier. Biological Reviews 100(1): 227–244. DOI: 10.1111/brv.13138 Dobrolyubova T.V. 1984a. Springtail assemblages in the alpine landscapes of the northwestern Caucasus. In: M.S. Ghilarov, N.M. Chernova (Eds.): Fauna and Ecology of Springtails. Moscow: Nauka. P. 89–95. [In Russian] Dobrolyubova T.V. 1984b. Springtail fauna of dark coniferous forests of the Northwest Caucasus. In: M.S. Ghilarov, N.M. Chernova (Eds.): Fauna and Ecology of Springtails. Moscow: Nauka. P. 95–98. [In Russian] Dobrolyubova T.V. 1987. Population structure and dynamics of springtail populations in mountain forest soils of the North-West Caucasus. PhD Thesis. Moscow: Moscow State Pedagogical Institute. 234 p. [In Russian] Dobrolyubova T.V. 1988. Population characteristics of springtails of mountain pine forests of Teberda. In: N.M. Chernova (Ed.): Ecology of Forest Soil Microarthropods. Moscow: Nauka. P. 60–65. [In Russian] Dobrolyubova T.V. 1995. Populations of Springtails of mountain pine forests at different altitudes (Caucasus). Ekologiya 2: 161–163. [In Russian] Engelmann H.D. 1978. Zur Dominanzklassifizierung von Bodenarthropoden. Pedobiologia 18(5–6): 378–380. DOI: 10.1016/S0031-4056(23)00612-1 Ficetola G.F., Marta S., Guerrieri A., Gobbi M., Ambrosini R., Fontaneto D., Zerboni A., Poulenard J., Caccianiga M., Thuiller W. 2021. Dynamics of Ecological Communities Following Current Retreat of Glaciers. Annual Review of Ecology, Evolution, and Systematics 52: 405–426. DOI: 10.1146/annurev-ecolsys-010521-040017 Gazaev Kh.M.M., Bozieva Zh.Ch. 2020. Comparison of the temperature in the surface layer of the atmosphere at two meteorological points in Kabardino-Balkaria State Nature Reserve. Bulletin of the Vladikavkaz Scientific Center 20(1): 62–66. DOI: 10.23671/VNC.2020.1.56949 [In Russian] Gazaev Kh.M.M., Agoeva E.A., Bozieva Zh.Ch., Ittiev A.B. 2018. Space-tempory changes of meteorological and hydrochemical parameters in the Bessengian Gorge. Physical Geography and Biogeography, Soil Geography and Landscape Geochemistry 4(71): 166–177. [In Russian] Gobbi M., Fontaneto D., De Bernardi F. 2006a. Influence of climate changes on animal communities in space and time: The case of spider assemblages along an alpine glacier foreland. Global Change Biology 12(10): 1985–1992. DOI: 10.1111/j.1365-2486.2006.01236.x Gobbi M., De Bernardi F., Pelfini M., Rossaro B., Brandmayr P. 2006b. Epigean arthropod succession along a 154-year glacier foreland chronosequence in the Forni Valley (Central Italian Alps). Arctic, Antarctic, and Alpine Research 38(3): 357–362. DOI: 10.1657/1523-0430(2006)38[357:EASAAY]2.0.CO;2 Gobbi M., Rossaro B., Vater A., De Bernardi F., Pelfini M., Brandmayr P. 2007. Environmental features influencing Carabid beetle (Coleoptera) assemblages along a recently deglaciated area in the Alpine region. Ecological Entomology 32(6): 682–689. DOI: 10.1111/j.1365-2311.2007.00912.x Gobbi M., Isaia M., De Bernardi F. 2011. Arthropod colonisation of a debris-covered glacier. Holocene 21(2): 343–349. DOI: 10.1177/0959683610374885 Gobbi M., Ballarin F., Brambilla M., Compostella C., Isaia M., Losapio G., Maffioletti C., Seppi R., Tampucci D., Caccianiga M. 2017. Life in harsh environments: Carabid and spider trait types and functional diversity on a debris-covered glacier and along its foreland. Ecological Entomology 42(6): 838–848. DOI: 10.1111/een.12456 Gravesen E., Dušátková L., Athey K.J., Qin J., Krogh P.H. 2024. Arthropod Food Webs in the Foreland of a Retreating Greenland Glacier: Integrating Molecular Gut Content Analysis With Structural Equation Modelling. Ecology and Evolution 14(12): e70687. DOI: 10.1002/ece3.70687 Greenslade P. 2007. The potential of Collembola to act as indicators of landscape stress in Australia. Australian Journal of Experimental Agriculture 47(4): 424–434. DOI: 10.1071/EA05264 Gwiazdowicz D.J., Zawieja B., Olejniczak I., Skubała P., Gdula A.K., Coulson S.J. 2020. Changing Microarthropod Communities in Front of a Receding Glacier in the High Arctic. Insects 11(4): 226. DOI: 10.3390/insects11040226 Hågvar S. 2010. Primary succession of springtails (Collembola) in a Norwegian glacier foreland. Arctic, Antarctic, and Alpine Research 42(4): 422–429. DOI: 10.1657/1938-4246-42.4.422 Hågvar S. 2012. Primary Succession in Glacier Forelands: How Small Animals Conquer New Land Around Melting Glaciers. In: S. Young (Ed.): International Perspectives on Global Environmental Change. Rijeka: InTech. P. 151–172. DOI: 10.5772/26536 Hågvar S. 2025. Primary succession near a melting glacier: How a fresh moraine developed into a complex ecosystem during eight years. Norwegian Journal of Entomology 72(1): 74–93. DOI: 10.61698/nje.v72n1.8 Hågvar S., Gobbi M. 2022. The role of arthropods in early colonization near melting glaciers: Contradictions between ecological assumptions and recent study results. Acta Oecologica 114: 103820. DOI: 10.1016/j.actao.2022.103820 Hågvar S., Pedersen A. 2015. Food Choice of Invertebrates During Early Glacier Foreland Succession. Arctic, Antarctic, and Alpine Research 47(3): 561–572. DOI: 10.1657/AAAR0014-046 Hågvar S., Solhøy T., Mong C.E. 2009. Primary Succession of Soil Mites (Acari) in a Norwegian Glacier Foreland, with Emphasis on Oribatid Species. Arctic, Antarctic, and Alpine Research 41(2): 219–227. DOI: 10.1657/1938-4246-41.2.219 Hågvar S., Gobbi M., Kaufmann R., Ingimarsdóttir M., Caccianiga M., Valle B., Pantini P., Fanciulli P.P., Vater A. 2020. Ecosystem Birth near Melting Glaciers: A Review on the Pioneer Role of Ground-Dwelling Arthropods. Insects 11(9): 644. DOI: 10.3390/insects11090644 Hågvar S., Valle B., Gobbi M. 2024. Remarks and advice to the study of early arthropod succession near melting glaciers. Arctic, Antarctic, and Alpine Research 56(1): 2335687. DOI: 10.1080/15230430.2024.2335687 Hammer Ø., Harper D.A.T., Ryan P.D. 2001. PAST: Paleontological statistics software package for education and data analysis. Palaeontologia Electronica 4(1): 9. Hock R., Bliss A., Marzeion B., Giesen R.H., Hirabayashi Y., Huss M., Radić V., Slangen A.B.A. 2019. GlacierMIP – A model intercomparison of global-scale glacier mass-balance models and projections. Journal of Glaciology 65(251): 453–467. DOI: 10.1017/jog.2019.22 Hodkinson I.D., Webb N.R., Coulson S.J. 2002. Primary community assembly on land – the missing stages: why are the heterotrophic organisms always there first?. Journal of Ecology 90(3): 569–577. DOI: 10.1046/j.1365-2745.2002.00696.x Hodkinson I.D., Coulson S.J., Webb N.R. 2004. Invertebrate community assembly along proglacial chronosequences in the high Arctic. Journal of Animal Ecology 73(3): 556–568. DOI: 10.1111/j.0021-8790.2004.00829.x Janetschek H. 1949. Tierische Successionen auf hochalpinem Neuland. Berichte des naturwissenschaftlich-medizinischen Vereins Innsbruck 48/49: 1–215. Janetschek H. 1958. Über die tierische Wiederbesiedlung im Hornkees-Vorfeld (Zillertaler Alpen). Schlern-Schriften 188: 209–246. Joosse E.N.G. 1965. Pitfall-trapping as a method for studying surface-dwelling Collembola. Zeitschrift für Morphologie und Ökologie der Tiere 55: 587–596. Kaprus' I.Ya. 2010. Macrogeographical trends of springtails (Collembola) taxonomical diversity. Scientific Bulletin of Uzhhorod University. Series Biology 28: 106–114. [In Ukrainian] Kaufmann R. 2001. Invertebrate succession on an Alpine glacier foreland. Ecology 82(8): 2261–2278. DOI: 10.1890/0012-9658(2001)082[2261:ISOAAG]2.0.CO;2 Kaufmann R., Fuchs M., Gosterxeier N. 2002. The soil fauna of an alpine glacier foreland: Colonization and succession. Arctic, Antarctic, and Alpine Research 34(3): 242–250. DOI: 10.1080/15230430.2002.12003491 Kawaue T., Nakamori T., Iwasaki Y., Potapov M. 2016. Comparison of sampling methods for Collembola on a cobble-dominated riverbank. Edaphologia 98: 21–27. DOI: 10.20695/edaphologia.98.0_21 König T., Kaufmann R., Scheu S. 2011. The formation of terrestrial food webs in glacier foreland: Evidence for the pivotal role of decomposer prey and intraguild predation. Pedobiologia 54(2): 147–152. DOI: 10.1016/j.pedobi.2010.12.004 Kushnov I., Abakumov E., Tembotov R., Nizamutdinov T. 2022. Migration of organic carbon and trace elements in the system glacier-soil in the Central Caucasus alpine environment. Journal of Mountain Science 19(12): 3458–3474. DOI: 10.1007/s11629-022-7589-x Kuznetsova N.A. 1985. Fauna and populations of springtails (Collembola) in coniferous forests of the European part of the USSR. PhD Thesis. Moscow. 286 p. [In Russian] Kuznetsova N.A. 2005. Organization of soil-dwelling springtail communities. Moscow: Prometey. 244 p. [In Russian] Kuznetsova N.A., Bokova A.I., Saraeva A.K., Shveenkova Yu.B. 2019. Structure of the species diversity of soil springtails (Hexapoda, Collembola) in pine forests of the Caucasus and the Russian plain: A multi-scale approach. Entomological Review 99(2): 143–157. DOI: 10.1134/S0013873819020027 Makarova O.L., Antipova M.D., Babenko A.B., Bushueva I.S., Chulei A.D., Golovatch S.I., Doroshina G.Y., Kolesnikov V.B., Mazei Y.A., Makarov K.V., Palatov D.M., Ponomarev A.V., Popov K.P., Rapoport I.B., Semionenkov O.I., Snegovaya N.Y., Sheftel B.I., Tsyganov A.N., Turbanov I.S., Zuev R.V. 2024. Successions of terrestrial invertebrate communities during the Tsey Glacier retreat, Central Caucasus. Caucasiana 3: 41–87. DOI: 10.3897/caucasiana.3.e117332 Mertens J., Beladjal L., Janssens F., Matthys P. 2007. Pitfall trapping in flooding habitats: a new technique reveals Archisotoma pulchella (Collembola: Isotomidae) as new to the Belgian fauna. Belgian Journal of Zoology 137: 177–181. Missa O., Basset Y., Alonso A., Miller S.E., Curletti G., De Meyer M., Eardley C., Mansell M.W., Wagner T. 2008. Monitoring arthropods in a tropical landscape: relative effects of sampling methods and habitat types on trap catches. Journal of Insect Conservation 13: 103–118. DOI: 10.1007/s10841-007-9130-5 Myers N., Mittermeier R.A., Mittermeier C.G., da Fonseca G.A., Kent J. 2000. Biodiversity hotspots for conservation priorities. Nature 403(6772): 853–858. DOI: 10.1038/35002501 Oksanen J., Simpson G., Blanchet F., Kindt R., Legendre P., Minchin P., O'Hara R., Solymos P., Stevens M., Szoecs E., Wagner H., Barbour M., Bedward M., Bolker B., Borcard D., Carvalho G., Chirico M., De Caceres M., Durand S., Evangelista H., FitzJohn R., Friendly M., Furneaux B., Hannigan G., Hill M., Lahti L., McGlinn D., Ouellette M., Ribeiro Cunha E., Smith T., Stier A., Ter Braak C., Weedon J., Borman T. 2025. vegan: Community Ecology Package. R package version 2.6-10. Available from https://CRAN.R-project.org/package=vegan Ponge J.F., Gillet S., Dubs F., Fedoroff E., Haese L., Sousa J.P., Lavelle P. 2003. Collembolan communities as bioindicators of land use intensification. Soil Biology and Biochemistry 35(6): 813–826. DOI: 10.1016/S0038-0717(03)00108-1 Potapov M. 2001. Synopses on Palaearctic Collembola. Vol. 3: Isotomidae. Görlitz: Naturkundemuseum. 603 p. Potapov M.B., Kuznetsova N.A. 2011. Methods for studying microarthropod communities: a manual for students and graduate students. Moscow: KMK Scientific Press Ltd. 89 p. [In Russian] Querner P., Bruckner A. 2010. Combining pitfall traps and soil samples to collect Collembola for site scale biodiversity assessments. Applied Soil Ecology 45(3): 293–297. DOI: 10.1016/j.apsoil.2010.05.005 R Core Team. 2024. R: A language and environment for statistical computing. Vienna: R Foundation for Statistical Computing. Available from https://www.R-project.org/ Raso L., Sint D., Mayer R., Plangg S., Recheis T., Brunner S., Kaufmann R., Traugott M. 2014. Intraguild predation in pioneer predator communities of alpine glacier forelands. Molecular Ecology 23(15): 3744–3754. DOI: 10.1111/mec.12649 Rounce D.R., Hock R., Maussion F., Hugonnet R., Kochtitzky W., Huss M., Berthier E., Brinkerhoff D., Compagno L., Copland L., Farinotti D., Menounos B., McNabb R.W. 2023. Global glacier change in the 21st century: Every increase in temperature matters. Science 379(6627): 78–83. DOI: 10.1126/science.abo1324 Schneider C., D'Haese C.A. 2023. Breakaway from a globular body shape: molecular phylogeny reveals the evolutionary history of the enigmatic springtail Mackenziella psocoides. Arthropod Systematics and Phylogeny 81: 781–799. DOI: 10.3897/asp.81.e104522 Seniczak A., Solhøy T., Seniczak S. 2006. Oribatid mites (Acari: Oribatida) in the glacier foreland at Hardangerjøkulen (Norway). Biological Letters 43(2): 231–235. Shveenkova Yu.B. 2022. The collembolan fauna (Hexapoda: Collembola) of The "Privolzhskaya Lesostep" Reserve and adjacent territories In the Penza Region. Russian Journal of Ecosystem Ecology 7(4). DOI: 10.21685/2500-0578-2022-4-4 [In Russian] Sint D., Kaufmann R., Mayer R., Traugott M. 2019. Resolving the predator first paradox: Arthropod predator food webs in pioneer sites of glacier forelands. Molecular Ecology 28(2): 336–347. DOI: 10.1111/mec.14839 Sokolov V.E., Tembotov A.K. 1989. Vertebrates of the Caucasus. Mammals. Insectivores. Moscow: Nauka. 548 p. [In Russian] Solomina O.N., Bradley R.S., Jomelli V., Geirsdottir A., Kaufman D.S., Koch J., McKay N.P., Masiokas M., Miller G., Nesje A., Nicolussi K., Owen L.A., Putnam A.E., Wanner H., Wiles G., Yang B. 2016. Glacier fluctuations during the past 2000 years. Quaternary Science Reviews 149: 61–90. DOI: 10.1016/j.quascirev.2016.04.008 Solomina O., Jomelli V., Bushueva I.S. 2024. Holocene glacier variations in the Northern Caucasus, Russia. In: European Glacial Landscapes. Amsterdam: Elsevier. P. 353–365. DOI: 10.1016/B978-0-323-99712-6.00005-2 Stebaeva S.K. 1970. Life forms of springtails (Collembola). Zoologicheskii Zhurnal 49(10): 1437–1454. [In Russian] Striuchkova A., Antipova M., Potapov M., Semenova D., Kuznetsova N. 2024. Genetic lineages of Parisotoma notabilis sensu lato (Collembola) in Eastern Europe and the Caucasus. Soil Organisms 96(1): 23–36. DOI: 10.25674/356 Taskaeva A.M. 2005. Springtail fauna of the Komi Republic. Bulletin of the Institute of Biology of Komi Scientific Center UB RAS 10(96): 16–21. [In Russian] Tielidze L.G., Wheate R.D. 2018. The greater Caucasus glacier inventory (Russia, Georgia and Azerbaijan). Cryosphere 12: 81–94. DOI: 10.5194/tc-12-81-2018 Tielidze L.G., Solomina O.N., Jomelli V., Dolgova E.A., Bushueva I.S., Mikhalenko V.N., Brauche R., ASTER T. 2020. Change of Chalaati Glacier (Georgian Caucasus) since the Little Ice Age based on dendrochronological and Beryllium‑10 data. Ice and Snow 60(3): 453–470. DOI: 10.31857/S2076673420030052 Tielidze L.G., Nosenko G.A., Khromova T.E., Paul F. 2022. Strong acceleration of glacier area loss in the Greater Caucasus between 2000 and 2020. Cryosphere 16: 489–504. DOI: 10.5194/tc-16-489-2022 Tielidze L.G., Mackintosh A.N., Gavashelishvili A., Gadrani L., Nadaraia A., Elashvili M. 2025. Post-Little Ice Age Equilibrium-Line Altitude and Temperature Changes in the Greater Caucasus Based on Small Glaciers. Remote Sensing 17(9): 1486. DOI: 10.3390/rs17091486 Tsepkova N.L. 2011. Meadow communities in Cherek-Bezengi canyon (Kabardino-Balkaria State High-Mountain Reserve). Proceedings of the Kabardino-Balkaria Scientific Centre of RAS 6: 57–64. [In Russian] Uligova T.S., Gedgafova F.V., Gorobtsova O.N., Tsepkova N.L., Rapoport I.B., Tembotov R.Kh., Khakunova E.M. 2019. Meadow biogeocenoses in the subalpine belt of the Kabardino-Balkaria State High-Mountain Reserve (Central Caucasus). Nature Conservation Research 4(2): 29–47. DOI: 10.24189/ncr.2019.012 [In Russian] Valle B., di Musciano M., Gobbi M., Bonelli M., Colonnelli E., Gardini G., Migliorini M., Pantini P., Zanetti A., Berrilli E., Frattaroli A.R., Fugazza D., Invernizzi A., Caccianiga M. 2022. Biodiversity and ecology of plants and arthropods on the last preserved glacier of the Apennines mountain chain (Italy). Holocene 32(8): 853–865. DOI: 10.1177/09596836221096292 Valle B., Gobbi M., Brambilla M., Borgatti M.S., Caccianiga M. 2023. Finding the optimal strategy for quantitative sampling of springtails community (Hexapoda: Collembola) in glacial lithosols. Pedobiologia 101: 150914. DOI: 10.1016/j.pedobi.2023.150914 Valle B., Barbon G., Cucini C., Nardi F., Ambrosini R., Boschi S., Buda J., Ficetola G.F., Frati F., Kováč Ľ., Marta S., Scotti R., Rivalta V.T., Zimmer A., Gobbi M., Caccianiga M. 2025. The Unexplored Biodiversity of 'Glacier Fleas' (Hexapoda: Collembola): Taxonomy, Distribution and Ecology in the European Alps and Apennines. Journal of Zoological Systematics and Evolutionary Research 2025: 1616350. DOI: 10.1155/jzs/1616350 Van Straalen N.M. 1997. Community structure of soil arthropods as bioindicator of soil health. In: C. Pankhurst, B.M. Doube, V.V.S.R. Gupta (Eds.): Biological indicators of soil health. Wallingford: CAB International. P. 235–264. Van Straalen N.M. 1998. Evaluation of bioindicator systems derived from soil arthropod communities. Applied Soil Ecology 9(1–3): 429–437. DOI: 10.1016/S0929-1393(98)00101-2 Walker L.R., Wardle D.A., Bardgett R.D., Clarkson B.D. 2010. The use of chronosequences in studies of ecological succession and soil development. Journal of Ecology 98(4): 725–736. DOI: 10.1111/j.1365-2745.2010.01664.x Wickham H. 2016. Data analysis. In: ggplot2. Use R!. Cham: Springer. P. 189–201. DOI: 10.1007/978-3-319-24277-4_9 Wickham H., Averick M., Bryan J., Chang W., McGowan L.D., François R., Grolemund G., Hayes A., Henry L., Hester J., Kuhn M., Pedersen T.L., Miller E., Bache S.M., Müller K., Ooms J., Robinson D., Seidel D.P., Spinu V., Takahashi K., Vaughan D., Wilke C., Woo K., Yutani H. 2019. Welcome to the tidyverse. Journal of Open Source Software 4(43): 1686. DOI: 10.21105/joss.01686 Zemp M., Frey H., Gärtner-Roer I., Nussbaumer S.U., Hölzle M., Paul F., Haeberli W., Denzinger F., Ahlstrøm A.P., Anderson B., Bajracharya S., Baroni C., Braun L.N., Cáceres B.E., Casassa G., Cobos G., Dávila L.R., Delgado G.H., Demuth M.N., Espizua L., Fischer A., Fujita K., Gadek B., Ghazanfar A., Hagen J.O., Holmlund P., Karimi N., Li Z., Pelto M., Pitte P., Popovnin V.V. et al. 2015. Historically unprecedented global glacier decline in the early 21st century. Journal of Glaciology 61(228): 745–762. DOI: 10.3189/2015JoG15J017 |