Articles | Volume 23, issue 12
https://doi.org/10.5194/bg-23-4199-2026
© Author(s) 2026. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
https://doi.org/10.5194/bg-23-4199-2026
© Author(s) 2026. This work is distributed under
the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Invertebrate communities on seasonal snow patches in the European lowlands are shaped by tree canopy-derived organic inputs
Dzmitry Lukashanets
CORRESPONDING AUTHOR
Marine Research Institute, Klaipėda University, Klaipėda, Lithuania
Krzysztof Zawierucha
Department of Animal Taxonomy and Ecology, Faculty of Biology, Adam Mickiewicz University, Poznań, Poland
Department of Antarctic Biology, Institute of Biochemistry and Biophysics, Polish Academy of Sciences, Warsaw, Poland
Jolita Petkuvienė
Marine Research Institute, Klaipėda University, Klaipėda, Lithuania
Donata Overlingė
Marine Research Institute, Klaipėda University, Klaipėda, Lithuania
Diana Vaičiūtė
Marine Research Institute, Klaipėda University, Klaipėda, Lithuania
Greta Kalvaitienė
Marine Research Institute, Klaipėda University, Klaipėda, Lithuania
Marija Kataržytė
Marine Research Institute, Klaipėda University, Klaipėda, Lithuania
Related authors
No articles found.
Sławomir Sułowicz, Krzysztof Zawierucha, Anna Markowicz, Krystyna Kozioł, Wiktoria Zientak, Adam Nawrot, Krzesimir Tomaszewski, Christoph Keuschnig, Bartłomiej Luks, and Catherine Larose
Biogeosciences, 23, 3023–3038, https://doi.org/10.5194/bg-23-3023-2026, https://doi.org/10.5194/bg-23-3023-2026, 2026
Short summary
Short summary
Our research shows that animals such as birds and mammals shaping the bacteria community structure found in Arctic snow. By analyzing snow samples from coastal Spitsbergen, we found that microbes linked to animal waste were common and influenced the types of bacteria present. This suggests that wildlife, not just wind or air, helps bring microbes into snow. Understanding this helps us better predict how Arctic ecosystems respond to environmental change and how life adapts in extreme conditions.
Tereza Novotná Jaroměřská, Roberto Ambrosini, Dorota Richter, Miroslawa Pietryka, Przemyslaw Niedzielski, Juliana Souza-Kasprzyk, Piotr Klimaszyk, Andrea Franzetti, Francesca Pittino, Lenka Vondrovicová, Tyler Kohler, and Krzysztof Zawierucha
Biogeosciences Discuss., https://doi.org/10.5194/bg-2022-198, https://doi.org/10.5194/bg-2022-198, 2022
Preprint withdrawn
Short summary
Short summary
Changes in the composition and biomass of photoautotrophs and consumers on glacier indicated phenological or ecological controls over their communities. We demonstrated that the recognition of the community structure of cryoconite holes requires a broad-scale and seasonal approach since biological communities vary in time and space on the glacier surface.
Cited articles
Anesio, A. M. and Laybourn-Parry, J.: Glaciers and ice sheets as a biome, Trends Ecol. Evol., 27, 219–225, 2012.
Bertolani, R., Rebecchi, L., and Beccaccioli, G.: Dispersal of Ramazzottius and other tardigrades in relation to type of reproduction, Invertebr. Reprod. Dev., 18, 153–157, https://doi.org/10.1080/07924259.1990.9672137, 1990.
Bosson, J. B., Huss, M., Cauvy-Fraunié, S., Clément, J. C., Costes, G., Fischer, M., Poulenard, J., and Arthaud, F.: Future emergence of new ecosystems caused by glacial retreat, Nature, 620, 562–569, https://doi.org/10.1038/s41586-023-06302-2, 2023.
Bouchard Jr., R. W., Schuetz, B. E., Ferrington Jr., L. C., and Kells, S. A.: Cold hardiness in the adults of two winter stonefly species: Allocapnia granulata (Claassen, 1924) and A. pygmaea (Burmeister, 1839) (Plecoptera: Capniidae), Aquat. Insect., 31, 145–155, 2009.
Brauer, F.: Insecten-Leben im Winter, Vereine zur Verbreitung naturwissenschaftlicher Kenntnisse im Wien, 357–381, 1871.
Chapman, J. A.: Observations on snow insects in western Montana, Can. Entomol., 86, 357–363, 1954.
Crosta, A., Valle, B., Caccianiga, M., Gobbi, M., Ficetola, F. G., Pittino, F., Franzetti, A., Azzoni, R. S., Lencioni, V., Senese, A., and Corlatti, L.: Ecological interactions in glacier environments: a review of studies on a model Alpine glacier, Biol. Rev., 100, 227–244, 2025.
Devetter, M., Renčo, M., Háněl, L., Jimenez-Santos, M. A., Doležálková, M. K., and Janko, K.: Rare presence of nematodes in cryoconite holes: evidence from a norwegian glacier, Hydrobiologia, 853, 865–879, https://doi.org/10.1007/s10750-025-05971-6, 2025.
Dial, C. R., Dial, R. J., Saunders, R., Lang, S. A., Lee, B., Wimberger, P., Dinapoli, M. S., Egiazarov, A. S., Gipple, S. L., Maghirang, M. R., Swartley-McArdle, D. J., Yudkovitz S. R., and Shain, D. H.: Historical biogeography of the North American glacier ice worm, Mesenchytraeus solifugus (Annelida: Oligochaeta: Enchytraeidae), Mol. Phylogenet. Evol., 63, 577–584, 2012.
Dimattia, L. M., Saglam, N., Saunders, R., and Shain, D. H.: An ice inhabiting bdelloid rotifer from North America, Extremophiles, 29, 30, https://doi.org/10.1007/s00792-025-01390-6, 2025.
Domine, F.: Should we not further study the impact of microbial activity on snow and polar atmospheric chemistry?, Microorganisms, 7, 260, https://doi.org/10.3390/microorganisms7080260, 2019.
Dong, Z., Jiang, H., Baccolo, G., Di Mauro, B., and Zawierucha, K.: Biological and pollution aerosols on snow and ice: interplay between the atmosphere and the cryosphere, J. Earth Sci., 34, 1951–1956, https://doi.org/10.1007/s12583-023-2004-2, 2023.
European Committee for Standardization: Water quality – Guidance standard on the enumeration of phytoplankton using inverted microscopy (Utermöhl technique), EN 15204:2006, https://repository.oceanbestpractices.org/server9/api/core/bitstreams/827e1cd6-84f9-4771-a27d-815c95aedeed/content (last access: 20 October 2025), 2006.
European Environment Agency: Global and European temperatures, https://www.eea.europa.eu/en/analysis/indicators/global-and-european-temperatures, last access: 15 November 2025.
Federation, W. E. and Aph Association: Standard methods for the examination of water and wastewater, American Public Health Association (APHA), Washington, DC, USA, 21 pp., 2005.
Ferree, M. A. and Shannon, R. D.: Evaluation of a second derivative UV/visible spectroscopy technique for nitrate and total nitrogen analysis of wastewater samples, Water Res., 35, 327–332, 2001.
Fiołka, M. J., Takeuchi, N., Sofińska-Chmiel, W., Mieszawska, S., and Treska, I.: Morphological and physicochemical diversity of snow algae from Alaska, Sci. Rep.-UK, 10, 19167, https://doi.org/10.1038/s41598-020-76215-x, 2020.
Fontaneto, D.: Long-distance passive dispersal in microscopic aquatic animals, Mov. Ecol., 7, https://doi.org/10.1186/s40462-019-0155-7, 2019.
Fontaneto, D., Ficetola, G. F., Ambrosini, R., and Ricci, C.: Patterns of diversity in microscopic animals: are they comparable to those in protists or in larger animals?, Global Ecol. Biogeogr., 15, 153–162, https://doi.org/10.1111/j.1466-822X.2006.00193.x, 2006.
Fontaneto, D., Herniou, E. A., Barraclough, T. G., and Ricci. C.: On the global distribution of microscopic animals: New worldwide data on bdelloid rotifers, Zool. Stud., 46, 336–346, 2007.
Giannetti, D., Giovannini, I., Massa, E., Schifani, E., Rebecchi, L., Guidetti, R., and Grasso, D. A.: Dispersion and new shelters offered by ants: myrmecophoresy of tardigrades, Front. Zool., 22, 30, https://doi.org/10.1186/s12983-025-00581-3, 2025.
Gobbi, M. and Lencioni, V.: Glacial Biodiversity: Lessons from Ground-dwelling and Aquatic, Glaciers and the polar environment, 143, https://doi.org/10.5772/intechopen.92826, 2020.
Hågvar, S.: A review of Fennoscandian arthropods living on and in snow, Eur. J. Entomol., 107, 281–298, https://doi.org/10.14411/eje.2010.036, 2010.
Hågvar, S. and Krzemińska, E.: Contribution to the winter phenology of Trichoceridae (Diptera) in snow-covered southern Norway, Studia dipterologica, 14, 271–283, 2008.
Hanzelová, M., Vido, J., Škvarenina, J., Nalevanková, P., and Perháčová, Z.: Microorganisms in summer snow patches in selected high mountain ranges of Slovakia, Biologia, 73, 1177–1186, https://doi.org/10.2478/s11756-018-0136-0, 2018.
Harrell Jr., F. E.: Package “Hmisc” (Version 4.7-1), https://cran.r-project.org/web/packages/Hmisc/Hmisc.pdf (last access: 1 October 2025), 2022.
HELCOM: Guidelines for monitoring of phytoplankton species composition, abundance and biomass, HELCOM, Helsinki, Finland, 22 pp., https://doi.org/10.25607/OBP-1822, 2021.
Hodson, A., Anesio, A. M., Tranter, M., Fountain, A., Osborn, M., Priscu, J., Laybourn-Parry, J., and Sattler, B.: Glacial ecosystems, Ecol. Monogr., 78, 41–67, 2008.
Hodson, A. J., Mumford, P. N., Kohler, J., and Wynn, P. M.: The High Arctic glacial ecosystem: new insights from nutrient budgets, Biogeochemistry, 72, 233–256, 2005.
Hoham, R. W. and Remias, D.: Snow and glacial algae: a review, J. Phycol., 56, 264–282, https://doi.org/10.1111/jpy.12967, 2020.
Hotaling, S. Shain, D. H., Lang, S. A., Bagley, R. K., Tronstad, L. M., Weisrock, D. W., and Kelley, J. L.: Long-distance dispersal, ice sheet dynamics and mountaintop isolation underlie the genetic structure of glacier ice worms, P. Roy. Soc., 286, 20190983, https://doi.org/P10.1098/rspb.2019.0983, 2019.
Hotaling, S., Wimberger, P. H., Kelley, J. L., and Watts, H.: Macroinvertebrates on glaciers, Ecology, 101, 1–3, 2020.
International Organisation for Standardisation: Water quality. Determination of ammonium. Part 1: Manual spectrometric method, ISO 7150-1:1984, https://www.iso.org/standard/18782.html (last access: 12 November 2025), 1984.
International Organisation for Standardisation: Soil quality – Determination of organic and total carbon after dry combustion (“elemental analysis”), ISO 10694:1995, https://www.iso.org/standard/18782.html (last access: 10 November 2025), 1995.
International Organisation for Standardisation: Determination of nitrite. Molecular adsorption spectrometric method, ISOEN 26777:1993, https://standards.iteh.ai/catalog/standards/ cen/27c3cc18-7b44-4c2c-bd04-52ff233b7828/en-26777-1993? srsltid=AfmBOopvR1UlMOn5Lbp35GYEn9yHSb1sgM6zMts ECZ8wicu9y8eApW8T (last access: 31 October 2025), 1993.
International Organisation for Standardisation: Water quality – Determination of phosphorus – Ammonium molybdate spectrometric method, EN ISO 6878:2004, https://www.iso.org/standard/36917.html (last access: 15 October 2025), 2004.
Jaroměřská, T. N., Ambrosini, R., Richter, D., Pietryka, M., Niedzielski, P., Souza-Kasprzyk, J., Klimaszyk, P., Franzetti, A., Pittino, F., Vondrovicová, L., and Senese, A.: Insights Into Cryoconite Community Dynamics on the Alpine Glacier Throughout the Ablation Season, Ecol. Evol., 15, e71064, https://doi.org/10.1002/ece3.71064, 2025.
Jeffrey, S. W. and Humphrey, G. F.: New Spectrophotometric Equations for Determining Chlorophylls a, b, c1 and c2 in Higher Plants, Algae and Natural Phytoplankton, Biochem. Physiol. Pfl., 167, 191–194, https://doi.org/10.1016/S0015-3796(17)30778-3, 1975.
Kivinen, S., Kaarlejärvi, E., Jylhä, K., and Räisänen, J.: Spatiotemporal distribution of threatened high-latitude snowbed and snow patch habitats in warming climate, Environ. Res. Lett., 7, 034024, https://doi.org/10.1088/1748-9326/7/3/034024, 2012.
Krasting, J. P., Broccoli, A. J., Dixon, K. W., and Lanzante, J. R.: Future changes in Northern Hemisphere snowfall, J. Climate, 26, 7813–7828, 2013.
Leya, T.: Snow algae: adaptation strategies to survive on snow and ice, in: Polyextremophiles: Life under multiple forms of stress, edited by: Seckbach, J., Oren, A., and Stan-Lotter, H., Springer, Dordrecht, Netherlands, 401–423, https://doi.org/10.1007/978-94-007-6488-0_17, 2013.
Morek, W., Surmacz, B., López-López, A., and Michalczyk, Ł.: “Everything is not everywhere”: Time-calibrated phylogeography of the genus Milnesium (Tardigrada), Mol. Ecol., 30, 3590–3609, 2021.
Murray, J.: Antarctic Rotifera, British Antarctic Expedition 1907–9, 41–65, https://doi.org/10.5962/bhl.title.22427, 1910.
Oksanen, J., Blanchet, J. G., Friendly, M., Kindt, R., Legendre, P., McGlinn, D., Minchin, P. R., O'Hara, R. B., Simpson, G. L., Solymos, P., Stevens, M. H. M., Szoecs, E., and Wagner, H.: Vegan: Community Ecology Package. R package version 2.5-7, https://CRAN.R-project.org/package=vegan (last access: 5 October 2025), 2020.
Olenina, I., Hajdu, S., Edler, L., Andersson, A., Wasmund, N., Busch, S., Gobel, J., Gromisz S., Huseby, S., Huttunen, M., Jaanus, A., Kokkonen, P., Jurgensone, I., and Niemkiewicz, E.: Biovolumes and size-classes of phytoplankton in the Baltic Sea, HELCOM Baltic Sea Environmental Proceedings, 106, 1–144, 2006.
Ono, M. and Takeuchi, N.: The diel vertical migration of microbes within snowpacks driven by solar radiation and nutrients, Arct. Antarct. Alp. Res., 57, 2460253, https://doi.org/10.1080/15230430.2025.2460253, 2025.
Ono, M., Takeuchi, N., and Zawierucha, K.: Snow algae blooms are beneficial for microinvertebrates assemblages (Tardigrada and Rotifera) on seasonal snow patches in Japan, Sci. Rep.-UK, 11, 5973, https://doi.org/10.1038/s41598-021-85462-5, 2021.
Ono, M., Takeuchi, N., and Zawierucha, K.: Description of a new species of Tardigrada Hypsibius nivalis sp. nov. and new phylogenetic line in Hypsibiidae from snow ecosystem in Japan, Sci. Rep.-UK, 12, 14995, https://doi.org/10.1038/s41598-022-19183-8, 2022.
Ono, M., Kobayashi, K., Seto, D., Konishi, F., Wada, K., Usuba, S., and Takeuchi, N.: Temporal and vertical changes in snow microbial communities during the melting season below canopy in Northern Japan, The Cryosphere, 19, 5983–5999, https://doi.org/10.5194/tc-19-5983-2025, 2025.
Pittino, F., Zawierucha, K., Poniecka, E., Buda, J., Rosatelli, A., Zordan, S., Azzoni, R. S., Diolaiuti, G., Ambrosini, R., and Franzetti, A.: Functional and taxonomic diversity of anaerobes in supraglacial microbial communities, Microbiology Spectrum, 11, e01004-22, https://doi.org/10.1128/spectrum.01004-22, 2023.
Procházková, L., Remias, D., Bilger, W., Křížková, H., Řezanka, T., and Nedbalová, L.: Cysts of the Snow Alga Chloromonas krienitzii (Chlorophyceae) Show Increased Tolerance to Ultraviolet Radiation and Elevated Visible Light, Front. Plant Sci., 11, 617250, https://doi.org/10.3389/fpls.2020.617250, 2020.
Procházková, L., Remias, D., Suzuki, H., Kociánová, M., and Nedbalová, L.: Chloromonas rubrosalmonea sp. nov. causes blooms of salmon-red snow due to high astaxanthin and low chlorophyll content, Bot. Lett., 171, 1–19, https://doi.org/10.1080/23818107.2023.2301608, 2024.
Ramazzotti, G. and Maucci, W.: Il Phylum Tardigrada, Mem. dell'Ist. Ital. di Idrobiol. Pallanza, 41, 1–1012, 1983.
Remias, D., Procházková, L., Nedbalová, L., Benning, L. G., and Lutz, S.: Novel insights in cryptic diversity of snow and glacier ice algae communities combining 18S rRNA gene and ITS2 amplicon sequencing, FEMS Microbiol. Ecol., 99, fiad134, https://doi.org/10.1093/femsec/fiad134, 2023.
Ricci, C.: Ecology of bdelloids: how to be successful, Hydrobiologia, 147, 117–127, https://doi.org/10.1007/BF00025734, 1987.
Robertson, M. W., Russo, N. J., McInnes, S. J., Goffinet, B., and Jiménez, J. E.: Potential dispersal of tardigrades by birds through endozoochory: evidence from Sub-Antarctic White-bellied Seedsnipe (Attagis malouinus), Polar. Biol., 43, 899–902, https://doi.org/10.1007/s00300-020-02680-9, 2020.
Rozwalak, P. Podkowa, P., Buda, J., Niedzielski, P., Kawecki, S., Ambrosini, R., Azzoni, R. S., Baccolo, G., Ceballos, J. L., Cook, J., and Di Mauro, B.: Cryoconite – From minerals and organic matter to bioengineered sediments on glacier's surfaces, Sci. Total Environ., 807, 150874, https://doi.org/10.1016/j.scitotenv.2021.150874, 2022.
Schill, R. O. (Ed.): Water bears: The biology of tardigrades, Springer Cham, Switzerland, 419 pp., ISBN 978-3-319-95701-2, 2018.
Segawa, T., Matsuzaki, R., Takeuchi, N., Akiyoshi, A., Navarro, F., Sugiyama, S., Yonezawa, T., and Mori, H.: Bipolar dispersal of red-snow algae, Nat. Commun., 9, 3094, https://doi.org/10.1038/s41467-018-05521-w, 2018.
Shain, D. H. Halldórsdóttir, K., Pálsson, F., Aðalgeirsdóttir, G., Gunnarsson, A., Jónsson, Þ., Lang, S. A., Pálsson, H. S., Steinþórssson, S., and Arnason, E.: Colonization of maritime glacier ice by bdelloid Rotifera, Mol. Phylogenet. Evol., 98, 280–287, 2016.
Shain, D. H., Novis, P. M., Cridge, A. G., Zawierucha, K., Geneva, A. J., and Dearden, P. K.: Five animal phyla in glacier ice reveal unprecedented biodiversity in New Zealand's southern Alps, Sci. Rep.-UK, 11, 3898, https://doi.org/10.1038/s41598-021-83256-3, 2021.
Shain, D. H., Rogozhina, I., Fontaneto, D., Nesje, A., Saglam, N., Bartlett, J., Zawierucha, K., Kielland, Ø. N., Dunshea, G., Arnason, E., and Rosvold, J.: Ice-inhabiting species of Bdelloidea Rotifera reveal a pre-Quaternary ancestry in the Arctic cryosphere, Biol. Letters, 20, 20230546, https://doi.org/10.1098/rsbl.2023.0546, 2024.
Shmakova, L. Malavin, S., Iakovenko, N., Vishnivetskaya, T., Shain, D., Plewka, M., and Rivkina, E.: A living bdelloid rotifer from 24 000-year-old Arctic permafrost, Curr. Biol., 31, R712–R713, 2021.
Soszyńska-Maj, A., Paasivirta, L., and Giłka, W.: Why on the snow? Winter emergence strategies of snow-active Chironomidae (Diptera) in Poland, Insect Sci., 23, 754–770, 2016.
Utermöhl, H.: Zur Vervollkommnung der quantitativen Phytoplankton-Methodik, Int. Assoc. Theor. Appl. Limnol., 9, 1–38, 1958.
Valle, B., Porco, D., Skarżyński, D., Frati, F., Caccianiga, M., Rodriguez-Prieto, A., Zeni, M., and Gobbi, M.: Alpine blooming of “snow fleas”: the importance of snow for Alpine springtails (Hexapoda: Collembola) ecology and biodiversity, Rend. Lincei-Sci. Fis., 35, 163–180, 2024.
Voigt, T., Füssel, H. M., Gärtner-Roer, I., Huggel, C., Marty, C., and Zemp, M.: Impacts of climate change on snow, ice, and permafrost in Europe: Observed trends, future projections, and socio-economic relevance, ETC/ACC Technical Paper, 13, 1–117, 2010.
Wickham, H.: Ggplot2: Elegant Graphics for Data Analysis. R package version 3.5.1, https://cran.r-project.org/web/packages/ggplot2/index.html (last access: 20 October 2025), 2016.
Yakimovich, K. M., Engstrom, C. B., and Quarmby, L. M.: Alpine snow algae microbiome diversity in the Coast Range of British Columbia, Front. Microbiol., 11, 1721, https://doi.org/10.3389/fmicb.2020.01721, 2020.
Zawierucha, K. and Shain, D. H.: Disappearing Kilimanjaro snow – Are we the last generation to explore equatorial glacier biodiversity?, Ecol. Evol., 9, 8911–8918, 2019.
Zawierucha, K., Kolicka, M., Takeuchi, N., and Kaczmarek, Ł.: What animals can live in cryoconite holes? A faunal review, J. Zool., 295, 159–169, 2015.
Zawierucha, K. Buda, J., Pietryka, M., Richter, D., Łokas, E., Lehmann-Konera, S., Makowska, N., and Bogdziewicz, M.: Snapshot of micro-animals and associated biotic and abiotic environmental variables on the edge of the south-west Greenland ice sheet, Limnology, 19, 141–150, 2018.
Zawierucha, K., Buda, J., Azzoni, R. S., Niśkiewicz, M., Franzetti, A., and Ambrosini, R.: Water bears dominated cryoconite hole ecosystems: densities, habitat preferences and physiological adaptations of Tardigrada on an alpine glacier, Aquat. Ecol., 53, 543–556, https://doi.org/10.1007/s10452-019-09707-2, 2019.
Zawierucha, K., Porazinska, D. L., Ficetola, G. F., Ambrosini, R., Baccolo, G., Buda, J., Ceballos, J. L., Devetter, M., Dial, R., Franzetti, A., and Fuglewicz, U.: A hole in the nematosphere: tardigrades and rotifers dominate the cryoconite hole environment, whereas nematodes are missing, J. Zool., 313, 18–36, 2021.
Zawierucha, K. Trzebny, A., Buda, J., Bagshaw, E., Franzetti, A., Dabert, M., and Ambrosini, R.: Trophic and symbiotic links between obligate-glacier water bears (Tardigrada) and cryoconite microorganisms, PLoS One, 17, e0262039, https://doi.org/10.1371/journal.pone.0262039, 2022.
Zawierucha, K., Kašparová, E. Š., McInnes, S., Buda, J., Ambrosini, R., Devetter, M., Ficetola, G. F., Franzetti, A., Takeuchi, N., Horna, P., and Jaroměřská, T. N.: Cryophilic Tardigrada have disjunct and bipolar distribution and establish long-term stable, low-density demes, Polar Biol., 46, 1011–1027, 2023a.
Zawierucha, K., Stec, D., Dearden, P. K., and Shain, D. H.: Two new tardigrade genera from New Zealand's Southern Alp glaciers display morphological stasis and parallel evolution, Mol. Phylogenet. Evol., 178, 107634, https://doi.org/10.1016/j.ympev.2022.107634, 2023b.
Short summary
Seasonal snow patches in lowland forests may look lifeless, but they host diverse microscopic animals. Studying 40 sites in the Baltic States, we found that rotifers, tardigrades and nematodes dominate these short-lived habitats, sometimes reaching very high numbers. Their presence depends strongly on surrounding trees, which supply organic matter to the snow. These overlooked cryospheric ecosystems deserve attention in a warming climate.
Seasonal snow patches in lowland forests may look lifeless, but they host diverse microscopic...
Altmetrics
Final-revised paper
Preprint