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Editor’s Note: This is the tenth post in Part V of the Visual Cultures of the Circumpolar North series edited by Isabelle Gapp and guest edited by Sarah Pickman.
Growing above the Arctic Circle requires some big C3 energy. Over the course of the Cenozoic period (Earth’s current geological era, lasting from roughly sixty-six million years ago to the present), plants’ serendipitous molecular changes have developed a complex set of morphological adaptations to cold climates, the predominant adaptation being what is known as the C3 or Calvin cycle: a metabolic pathway whereby plants and algae convert inorganic carbon dioxide from the atmosphere into organic compounds. This process produces sugars that allow the plants to live (Figure 1). The main advantage of C3 photosynthesis in polar environments is its higher efficiency at low temperatures. Arctic plants’ ability to photosynthesize compounds from the atmosphere under continuous daylight during the long polar day, and a twenty-four hour assimilation mechanism for absorbing these materials, are some of their key ecophysiological characteristics.1 Though many plants have survived in the far North for thousands of years, their existence still astonished generations of outside travelers who viewed them.
Claiming space within the tundra, Arctic flora challenged Southerners’ perception that the Arctic was a world of white ice.2 These vegetal beings, over three thousand species and counting,3 respond to temperature fluctuations over the year. Their greens and browns signal moments of annual productivity and mortality that paint the landscape in saturated colors. Since the early modern period, outside visitors, including scientists, have collected northern plant specimens and preserved them in archives, museums, and university collections (Figure 2). They have formed key parts of many natural history collections around the globe.
Today, Arctic plant species are used as sources of data to study how biodiversity is responding to global climate changes, including understanding the impact of global warming on plant phenology, or the study of cyclical events in the natural world.4 Both contemporary and historic collections of Arctic flora are being deployed for this task. Utilizing this flora for scientific research requires using plant collections in ways that have exceeded their intended, instrumental functions of describing or categorizing. Researchers have reconfigured the traditional use of natural history collections for taxonomic, systematic, and biogeographical study into indicators of global climate change (Figure 3).5
Figure 3: From left to right: (1) a) Arctic willow (Salix arctica) herbarium specimen image collected in Greenland in 1908, with inset (b) showing septa (terminal bud scars) used to measure annual growth increments. On this specimen image, multiple years of growth are detectable providing growth increment data dating back several years. (2) Model estimated trends in growth over time and in response to climate from generalised additive models (GAMs) for each Salix species, plotted as the partial effects of (a) growth year and (b) temperature. Shaded ribbons around the fitted lines represent the 95% confidence intervals around the predicted mean growth values from GAMs. Figure and modified figure captions from Ahlstrand, Natalie Iwanycki, Zoe A. Panchen, Anne D. Bjorkman, and James DM Speed. “Herbarium specimens reveal drivers of Arctic shrub growth.” New Phytologist, Letter, 2025.
Yet the phenomenon of having cold adaptive traits is not going to assist Arctic species in tolerating forecasted warming trends, leading to increased plant mortality or landscape browning as observed at large geographic scales.6 Therefore, paradoxically the transformation of dried plant material in natural history collections into scatter graphs and maps created by contemporary researchers not only offers a new window into seeing climate change over time, but paints a new virtual and digital Arctic visual in computer-generated colors (Figure 4).
Figure 4: From left to right: (1) Location of sites of collected dwarf shrub samples from herbaria in Kew and Copenhagen. (2) Examples of cross sections of Salix arctica obtained from historical collections, (A. 1948, B. 1831, C. 1927,D. 1957), (3) Positions of growth ring sequences of arctic willow from NE Greenland, Thule, dated by the date on the sheet’s labelling and confirmed by dendrochronological cross-dating of chronologies. Figure and modified igure captions from Opała-Owczarek, Magdalena, and Piotr Owczarek. “Herbarium records in Arctic dwarf shrub dendrochronology: Methodological approach and perspectives.” Dendrochronologia 80 (2023): 126102.
The Arctic has drawn naturalists and botanists, particularly from places close to the tundra such as Norway, since the early nineteenth century.7 By moving through the networks of commercial, colonial, missionary, and scientific institutions, as well as Indigenous communities, collectors removed selected plants from their habitats to represent the diversity of northern flora. Arctic specimens were housed on shelves in museums and herbaria, transforming these organic forms into scientific objects (Figure 5).8 With the development of glacial science, Arctic plants, situated at the edge of the ice, challenged existing evolutionary and geological theories. By the mid-nineteenth century, the field of glacial studies argued that geological changes in northern environments had occurred due to vast ice sheets that had moved across them, scouring and breaking the land into the formations scientists now saw. If ice sheets decimated plant communities in the deep past, how were living species able to reemerge from such destruction while lacking powers of mobility and locomotion? The physical sciences could not answer this question without thinking with plants.
(licensed under http://creativecommons.org/licenses/by-sa/4.0/)
Beginning in 1820, collections of fossil plants from the Arctic circle placed paleobotany and phytogeography in the center of debates about evolutionary and climate history, and geology (Figure 6). Plant biogeography, the study of plant distribution which connected the organic and inorganic, offered a means to understand how all species formed assemblages and communities, and diverged from each other across historic, recent, and deep geological time (Figure 7).
Figure 6: From left to right: (1) Photographs from the collections. (A, C, D) Type collection showing storage of fossils together with the publication in which they were introduced, sorted chronologically according to year of publication. (B) General Arctic collection, sorted geographically and stratigraphically. (2) Fossils collected by Baltazar M. Keilhau during his 1827 expedition to Svalbard. (A) The Permian bryozoan Dyscritella from the Miseryfjellet Formation on Bjørnøya, PMO 245.673. (B) Historical label accompanying the fossil figured in A. (C) Permian spiriferid brachiopod from Sørkappland (“SydCap”), southern Spitsbergen, PMO 231.567. (D) A Holocene gastropod from Edgeøya (“Stans Land”), PMO 243.426. (E) Historical label accompanying the fossil figured in D. (F) An equisetacean specimen from the Carboniferous of Bjørnøya (“Bären Eiland”), PMO 170.014. Images taken from Nakrem, H. A., Lindemann, F.-J., Hurum, J. H., & Hammer, Ø. (2023). Fossils From the Arctic in the Collections of the Natural History Museum in Oslo, Norway. Collections: A Journal for Museum and Archives Professionals, 19(3), 420-441.
Figure 7: Comparative distributions of Arctic flora.Image taken from Markley, Paul T., Collin P. Gross, and Barnabas H. Daru. “The changing biodiversity of the Arctic flora in the Anthropocene.” American Journal of Botany 112, no. 2 (2025): e16466.
Historian of science Trevor Levere showed that it was circumpolar flora, more than the geological science of magnetism, that offered nineteenth-century scientists a key methodological approach to understand the tundra. According to Levere plants “provided so powerful a motive for ever-extended arctic exploration,” ultimately enabling botanical scientists and explorers to develop models of integrated circumpolar sciences.9 Plants bridged the space between evolution, geology, and ecosystem engineers, transforming landscapes ever so slowly to suit their own needs. With their interest in integrated Arctic sciences, locating a suitable plant to reveal the Arctic’s past, present and future did not take botanists very long.
Dryas octopetala (known to Inuit as malikkaat), or mountain avens, is a mat-forming species that helps stabilize the ground for other plants to move in (Figure 8).10 With its heliotropic (tendency to grow towards the sun) and hygroscopic (meaning it readily absorbs surrounding water) qualities, Inuit relied on the plant as their co-companion; an ecological kinship. They ‘read’ the plant for a sense of the time of day, predicting weather changes, the season of the year, and even as “indicators of direction” as malikkaat follows the sun even in fog.11 This was a knowledge system carried generationally and tacitly learnt through movement. As one of the hardiest plants on the planet, with a robust dispersal system reliant on wind and ice across great distances, this long-lived plant (up to one hundred years) offered a means to understand climate change beyond human lifespans and into the historic and deep time. That scientists located numerous Dryas fossils from the deep past was a key development in trying to understand planetary-scale changes in Earth’s climate.
Figure 8: Dryas integrifolia. Photographed in Upernavik, Greenland 2007 by Kim Hansen. CC BY-SA 3.0 http://creativecommons.org/licenses/by-sa/3.0/, via Wikimedia Commons
In 2007 the Younger Dryas (YD) hypothesis emerged and gained traction within the scientific community. It was so named after the mountain avens because of the species’s abundant fossils in sediments (Figure 7).12 The Younger Dryas hypothesis suggested that a cometary or meteoritic body or bodies hit and/or exploded over North America around 12,900 years ago, causing the YD climate episode (a sudden abrupt cooling across the Earth’s northern hemisphere), the extinction of Pleistocene megafauna, the demise of the widespread Clovis archeological culture, and a range of other large-scale climatic shifts and knock-on effects on humans and animals. The warming that had begun to usher in the Holocene era at approximately 12,000 BCE ended abruptly with sudden cooling for 1,300 years after this sudden event, followed by rapid warming, with Greenland’s temperature increasing 15oF in less than a decade. Palynological (pollen) research revealed that alongside these changes, mountain avens migrated rapidly as the ice alternatively retreated and expanded.13 Today the Younger Dryas is the subject of intense scientific attention because it is the closest such rapid climate shift to our own Anthropocene, however, with systematic review of the evidence presented by the hypothesis and framing it as a “requiem,” suggesting the end of the YD hypothesis.14 The Dryas plant, both contemporary examples in situ and historic examples in herbaria, became a new kind of proxy of planetary climate change today.
Yet this proxy or sentinel for change would merely highlight that although there has been considerable effort in documenting the effects of climate on Arctic plants, there remained a gap in the understanding of long‐term effects of climate on Arctic vegetation in relation to its past, present, and future. What little data exists on Arctic plant species reports conflicting responses of plants to warming.15 The Dryas proxy would become a harbinger of the lack of scientific ability to truly predict what was going to happen on a planetary scale. Just as historic Arctic explorers and travelers were astonished by the unfamiliar scales of life, time, size, and space in the Arctic, the vast dimensions necessary for understanding climate change were amplified by the limitations of what could be comprehended today. This presented scientists and observers with a disorientating and intoxicating conundrum equal to encountering the vastness of the floral-scapes of the Arctic.
This, however, is not the only issue that studying Arctic flora like mountain avens presents. Collections of Arctic plants, both those collected recently and those assembled decades or centuries ago, are not comprehensive of all northern plant life. Plant collections contain biases, over- or under-representations, and gaps associated with collectors, seasonality, taxonomic biases, and geographic sampling localities (Figure 9).16
Figure 9: A heat map of the herbarium specimens collected across Nunavut (n = 26,558).Communities near higher record densities are labeled. Inset: map of Canada with Territory of Nunavut highlighted in black. Figure and modified figure caption from Panchen, Z. A., J. Doubt, H. M. Kharouba,and M. O. Johnston. 2019. Patterns and biases in an Arctic herbarium specimen collection: Implications for phenological research. Applications in Plant Sciences 7(3): e1229.
These biases become more pronounced in Arctic regions where collecting trips are under greater influence of funding, access, climate, and weather.17 As scientists continue to fill in the gaps in human knowledge using Arctic species as indicators, the proxies we rely on to tell us climate change is occurring are ironically the very lifeforms that are running out of time. Even with their big C3 energy many Arctic species are entering the elevator towards their own extinction.
Notes
[1] The C3 photosynthetic phenomenon was first described in 1928 by D. Müller in willow (Salix glauca) of western Greenland (“Die Kohlensäuressimilation bei Arktischen Pflanzen und die Abhängigkeit der Assimilation von der Temperatur,” Zeitschrift für wissenschaftliche Biologie. Abteilung E. Planta, Vol. 6, No. 1 (1928), pp. 22-39). Plant scientists confirmed Müller’s description of the mechanism (Sergeĭ Pavlovich Kostychev in 1930 on the Barents Sea coast and H. G. Wager in 1941 in eastern Greenland). Subsequent studies elucidated that light was assimilated continuously over the course of twenty-four hours making C3 photosynthesis more efficient per unit area at lower leaf temperatures. See: Alexander, Vera. “Arctic Ecosystems in a Changing Climate: An Ecophysiological Perspective.” BioScience 42, no. 9 (1992): 710-712; Semikhatova, O. A. “Photosynthesis, respiration, and growth of plants in the Soviet Arctic.” Arctic Ecosystems in a Changing Environment (1992): 169-192; Semikhatova, O.A., Ivanova, T.I. & Kirpichnikova, O.V. Comparative study of dark respiration in plants inhabiting arctic (Wrangel Island) and temperate climate zones. Russian Journal of Plant Physiology, 54, 582–588 (2007); Chapin III, F. Stuart, Robert L. Jefferies, James F. Reynolds, Gaius R. Shaver, Josef Svoboda, and Ellen W. Chu, eds. Arctic ecosystems in a changing climate: an ecophysiological perspective. Academic Press, 2012.Nickelsen, Kärin. Explaining Photosynthesis : Models of Biochemical Mechanisms, 1840-1960. Springer, 2015; Markovskaya, E.F. “Eco-physiological features of vascular plants in the arctic tundra of Western Spitsbergen.” Transactions of the Kola Science Centre of the Russian Academy of Sciences. Applied Ecology of the North. 2021, 12, 175–180; Hobbie, John, Gaius Shaver, Toke Thomas Høye, and Joseph Bowden. “Arctic Tundra.” In Arctic Ecology, edited by David N. Thomas, 103–30. Hoboken, NJ: Wiley, 2021; Hüner, Norman PA, David R. Smith, Marina Cvetkovska, Xi Zhang, Alexander G. Ivanov, Beth Szyszka-Mroz, Isha Kalra, and Rachael Morgan-Kiss. “Photosynthetic adaptation to polar life: Energy balance, photoprotection and genetic redundancy.” Journal of Plant Physiology 268 (2022): 153557;Vasilevskaya, N.V. Arctic Plants Under Environmental Stress: A Review. Stresses, 2025, 5(4), p.64.
[2] Danell, Kjell. “What Is the Arctic?” In Arctic Ecology, edited by David N. Thomas, 1–22. Hoboken, NJ: Wiley-Blackwell, 2021.
[3] About 2130 angiosperm species and subspecies occur in the Arctic and 84 species are endemic to the region with an estimated 900 species of Arctic bryophytes (mosses and liverworts). See: Daniëls, F., Gillespie, L. & Poulin, M. In Arctic Biodiversity Assessment: Status and Trends in Arctic Biodiversity (ed Meltofte, H.) 311–353. (Conservation of Arctic Flora and Fauna, Arctic Council, 2013); Aronsson, Mora, Daniëls, Fred J.A. Gillespie, Lynn, Heiðmarsson, Starri, Kristinsson, Hörður, M. Ickert-Bond, Stefanie, Väre, Henry and Westergaard, Kristine Bakke. Red Listing of Arctic Vascular Plants: Current Status and Recommendations. https://www.researchgate.net/publication/256548700_Red_Listing_of_Arctic_Vascular_Plants_Current_Status_and_Recommendations#fullTextFileContent[accessed Nov 23 2025].
[4] Heberling, J.M. “Herbaria as big data sources of plant traits.” International Journal of Plant Sciences, 2022, 183: 87–118; Meineke EK, Davies TJ, Daru BH, Davis CC. “Biological collections for understanding biodiversity in the Anthropocene.” Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences (2019), 374: 20170386; Meineke EK, Davis CC, Davies TJ. “The unrealized potential of herbaria for global change in biology.” Ecological Monographs (2018), 88: 505–525; Menzel A, Sparks TH, Estrella N, Koch E, Aasa A, Ahas R, Alm-Kubler K, Bissolli, P, Braslavska O, Briede A et al.” European phenological response to climate change matches the warming pattern.” Global Change Biology (2006) 12: 1969–1976; Lang PLM, Willems FM, Scheepens JF, Burbano HA, Bossdorf O. “Using herbaria to study global environmental change.” New Phytologist (2019) 221: 110–122; Mandrioli M.” From dormant collections to repositories for the study of habitat changes: the importance of herbaria in modern life sciences.” Life (2023) 13: 2310; Heberling JM, Isaac BL. “Herbarium specimens as exaptations: new uses for old collections.” American Journal of Botany (2017) 104: 963–965.
[5] Lane, MA. “Roles of natural history collections.” Annals of the Missouri Botanical Garden (1996) 83: 536–545; Epstein, H.E., Walker, D.A., Raynolds, M.K., Jia, G.J., Kelley, A.M. “Phytomass patterns across a temperature gradient of the north American arctic tundra.” Journal of Geophysical Research (2018) 113, G03S02; Epstein, H. E., D. A. Walker, G. V. Frost, M. K. Raynolds, U. Bhatt, R. Daanen, B. Forbes, J. Geml, E. Kaärlejarvi, O. Khitun, A. Khomutov, P. Kuss, M. Leibman, G. Matyshak, N. Moskalenko, P. Orekhov, V. E. Romanovsky and I. Timling “Spatial patterns of arctic tundra vegetation properties on different soils along the Eurasia Arctic Transect, and insights for a changing Arctic.” Environmental research letters, (2020) 16: 014008.
[6] Markley, Paul T., Collin P. Gross, and Barnabas H. Daru. “The changing biodiversity of the Arctic flora in the Anthropocene.” American Journal of Botany 112, no. 2 (2025): e16466.Crawford, R. M. Cold climate plants in a warmer world. Plant Ecology & Diversity (2008) 1: 285‐297; Callaghan, T. V., L. O. Björn, Y. Chernov, T. Chapin, T. R. Christensen, B. Huntley, R. A. Ims, et al. Synthesis of effects in four Arctic subregions. AMBIO: A Journal of the Human Environment (2004) 33: 469‐473; Bokhorst, S., L. Jaakola, K. Karppinen, G. K. Edvinsen, H. K. Mæhre, and J. W. Bjerke. “Contrasting survival and physiological responses of sub‐Arctic plant types to extreme winter warming and nitrogen.” Planta (2018) 247: 635‐648; Forrest J, Miller-Rushing AJ. “Toward a synthetic understanding of the role of phenology in ecology and evolution.” Philosophical Transactions of the Royal Society B. (2010) Oct 12;365(1555):3101-12; Raynolds, M. K., J. C. Comiso, D. A. Walker, and D. Verbyla.”Relationship between satellite‐derived land surface temperatures, arctic vegetation types, and NDVI.” Remote Sensing of Environment (2008) 112: 1884‐1894; Raynolds, M. K., D. A. Walker, A. Balser, C. Bay, M. Campbell, M. M. Cherosov, F. J. Daniëls, et al. “A raster version of the Circumpolar Arctic Vegetation Map (CAVM).” Remote Sensing of Environment (2019) 232: 111297; Henry, G. H., R. D. Hollister, K. Klanderud, R. G. Björk, A. D. Bjorkman, C. Elphinstone, I. S. Jónsdóttir, et al. “The International Tundra Experiment (ITEX): 30 years of research on tundra ecosystems.” Arctic Science, (2022) 8: 550‐571.
[7] One of the first explorers of the Arctic was the county commissioner Hans Hansen Lilienskiold (c. 1650–1703) who made a survey of the plant vegetation in Arctic parts of the Norwegian mainland already in the seventeenth century. See: Bjorå, Charlotte Sletten, Mika Bendiksby, Bjørn Petter Løfall, Lars Erik Johannesen, and Einar Timdal. “Collections of Arctic Plants, Lichens, and Fungi in the Natural History Museum, University of Oslo, Norway.” Collections 19, no. 3 (2023): 293-309; Iwanycki Ahlstrand, Natalie. “Digitization of the greenland vascular plant herbarium as a unique research infrastructure to study arctic climate change and inform nature management.” Collections 19, no. 3 (2023): 310-321.
[8] Craciun, Adriana. “The Flowers Opened the Way: Arctic Floras in Green Lands.” Environmental Humanities 17, no. 1 (2025): 129-153; Brochmann, Christian, Cassandra Elphinstone, Siri Birkeland, Hajime Ikeda, Pernille B. Eidesen, Inger G. Alsos & Kristine B. Westergaard, “Phylogeography of Arctic plants: where are we after 35 years, and where to go?” Plant Ecology & Diversity (29 Oct 2025).
[9] Levere, Trevor. Science in the Canadian Arctic: A Century of Exploration, 1818–1918. Cambridge: Cambridge University Press, 1993, p. 238.
[10] Welker, J.M., Molau, U., Parsons, A.N., Robinson, C.H. and Wookey, P.A. “Responses of Dryas octopetala to ITEX environmental manipulations: a synthesis with circumpolar comparisons.” Global Change Biology, (1997) 3: 61-73.
[11] Ziegler, Anna, Aalasi Joamie, and Rebecca Hainnu. Edible and Medicinal Arctic Plants: An Inuit Elder’s Perspective. 2nd ed. Iqaluit, NU: Inhabit Media, 2018; Cuerrier, Alain, Ashleigh Downing, Jill Johnstone, Luise Hermanutz, Laura Siegwart Collier, and Elders and Youth Participants of Nain and Old Crow. “Our Plants, Our Land: Bridging Aboriginal Generations through Cross-cultural Plant Workshops.” Polar Geography 35, nos. 3–4 (2012): 195–210; Haramincic, Arijana. “The Medicine Story: Secret Life of the Arctic Willow.” Turtle Island Journal of Indigenous Health (2023) 1, no. 3.
[12] Pinter, Nicholas, Andrew C. Scott, Tyrone L. Daulton, Andrew Podoll, Christian Koeberl, R. Scott Anderson, and Scott E. Ishman. “The Younger Dryas impact hypothesis: A requiem.” Earth-Science Reviews 106, no. 3-4 (2011): 247-264.
[13] Fiedel, Stuart J. “The Mysterious Onset of the Younger Dryas.” Quaternary International 242, no. 2 (2011): 262–66; Mangerud, Jan. “The Discovery of the Younger Dryas.” Boreas 50, no. 1 (2020): 1–5
[14] Pinter, Nicholas, Andrew C. Scott, Tyrone L. Daulton, Andrew Podoll, Christian Koeberl, R. Scott Anderson, and Scott E. Ishman. “The Younger Dryas impact hypothesis: A requiem.” Earth-Science Reviews 106, no. 3-4 (2011): 247-264.
[15] Huettmann, F. and S. M. Ickert‐Bond. “On open access, data mining and plant conservation in the Circumpolar North with an online data example of the Herbarium,” University of Alaska Museum of the North. Arctic Science, (2017) 4: 433‐470; Diepstraten, R. A., T. D. Jessen, C. M. Fauvelle, and M. M. Musiani. 2018. “Does climate change and plant phenology research neglect the Arctic tundra?” Ecosphere, 9: e02362; Bjorkman, A. D., M. García Criado, I. H. Myers‐Smith, V. Ravolainen, I. S. Jónsdóttir, K. B. Westergaard, J. P. Lawler, et al. “Status and trends in Arctic vegetation: Evidence from experimental warming and long‐term monitoring.” Ambio (2020) 49: 678‐69; Markley, Paul T., Collin P. Gross, and Barnabas H. Daru. “The changing biodiversity of the Arctic flora in the Anthropocene.” American Journal of Botany 112, no. 2 (2025): e16466.
[16] Daru, Barnabas H., Daniel S. Park, Richard B. Primack, Charles G. Willis, David S. Barrington,Timothy J.S. Whitfeld, Tristram G. Seidler, et al.. “Widespread Sampling Biases in Herbaria Revealed From Large-Scale Digitization.” New Phytologist (2018) 217 (2): 939–55; Ahlstrand, Natalie Iwanycki, Zoe A. Panchen, Anne D. Bjorkman, and James DM Speed. “Herbarium specimens reveal drivers of Arctic shrub growth.” New Phytologist (2025); Ahlstrand, Natalie Iwanycki, Richard B. Primack, Matthew W. Austin, Zoe A. Panchen, Christine Römermann, and Abraham J. Miller‐Rushing. “The promise of digital herbarium specimens in large‐scale phenology research.” New Phytologist (2025).
[17] Huettmann, F. and S. M. Ickert‐Bond. “On open access, data mining and plant conservation in the Circumpolar North with an online data example of the Herbarium,” University of Alaska Museum of the North. Arctic Science, (2017). 4: 433‐470; Panchen, Zoe A., Jennifer Doubt, Heather M. Kharouba, and Mark O. Johnston. “Patterns and Biases in an Arctic Herbarium Specimen Collection: Implications for Phenological Research.” Applications in Plant Sciences (2019) 7 (3).
Feature image: Mari Karlstad, Norges arktiske universitetsmusem. Image from “Botany: Botanical and Mycological Collections at the Arctic University Museum of Norway (TROM),” 2025, Scientific collection-based taxonomy and biosystematics (NatSciCol).
Nuala Caomhanach
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