Small but mighty: structures created by small mammals affect C and nutrient cycling in arctic tundra

While typically thought of as having top-down roles in ecosystems, small mammal herbivores bottom-up roles are often underappreciated. As structure builders, small mammals can impact ecosystem function through alterations of soil nutrient cycling. We examined the effects of small mammal-built structure types (hay piles, runways, latrines) on soil and plant biogeochemical cycling across three tundra ecosystems in northern Alaska.

We found that structures play important roles in regulating soil nitrogen levels, regardless of tundra ecosystem. However, different structures influence soil nutrients in different ways. Hay piles increased soil N and plant P, while latrines influenced soil C, N, and P, and runways mainly affected soil N. We also found strong changes in the cover of structures on tundra during different phases of the small mammal population cycle.

Structures built by small mammals have the ability to increase or decrease biogeochemical cycling rates and arctic ecosystem function. Additionally, changes in the abundance of these structures may create pulses of resource availability during the high phase of the population cycle or may aid in maintaining ecosystem function when animals are rare on the landscape. Our work highlights the roles that small mammals play in the bottom-up regulation of tundra ecosystems and the need to include small mammals in our understanding of ecosystem function.   

You can read the full paper here: https://doi.org/10.1111/1365-2435.14127

Reference: Roy, A., Gough, L., Boelman, N.T., Rowe, R.J., Griffin, K.L. and McLaren, J.R., 2022. Small but mighty: Impacts of rodent‐herbivore structures on carbon and nutrient cycling in arctic tundra. Functional Ecology.


Text: Austin Roy, University of Texas at El Paso. This paper is a contribution of Team Vole, a group of collaborative scientists seeking to understand the impacts of small herbivores on carbon and nutrient cycling in tundra ecosystems.

Photo credit: Luke Johnson

Herbivore species co-existence in changing rangeland ecosystems

Northern rangelands are changing fast, and these changes can have profound consequences to species coexistence and management.

In their recent paper, Noémie Boulanger-Lapointe and collaborators compiled occurrence data for the main vertebrate herbivore species present in the highlands of Iceland (sheep, reindeer, pink-footed goose and rock ptarmigan). They used an ensemble model workflow to analyse their distribution and its drivers and produced the first high-resolution national open-source and open-access models for Iceland.

The analyses show that vegetation productivity and soil type were the main drivers of herbivore species diversity across Iceland. The overlapping distributions of sheep and geese point out the potential for wildlife-livestock conflicts and for continued ecosystem degradation at higher elevations even under declining livestock abundance.

Compiling data on herbivore occurrence across Iceland was not a trivial task. Data sources ranged from GPS collar data to citizen science observations and span a long period of time (1861-2021). The data compilation provides the most extensive database on herbivore distribution in Iceland. Together with the open-access modelling workflow, such dataset provides a framework for transparent and repeatable science-based management decisions.

You can read the full paper here: https://www.sciencedirect.com/science/article/pii/S0048969722042371

Reference: Boulanger-Lapointe, N., Ágústsdóttir, K., Barrio, I.C., Defourneaux, M., Finnsdóttir, R., Jónsdóttir, I.S., Marteinsdóttir, B., Mitchell, C., Möller, M., Nielsen, Ó.K. and Sigfússon, A.Þ., 2022. Herbivore species coexistence in changing rangeland ecosystems: First high resolution national open-source and open-access ensemble models for Iceland. Science of The Total Environment, p.157140.


This paper was part of Noémie’s postdoc project funded by the University of Iceland and is a contribution to the TUNDRAsalad project

Photograph: Sheep and pink-footed geese in Iceland (photo: Maite Gartzia)

Growth rings show limited evidence for ungulates’ potential to suppress shrubs across the Arctic

Herbivores may counteract climate warming impacts on tundra by reducing plant growth. However, the strength of this effect may depend on prevailing climatic conditions. To study how ungulates interact with temperature to influence growth of tundra shrubs across the Arctic tundra biome, we assembled dendroecological data from 20 sites, comprising 1,153 individual shrubs and 22,363 annual growth rings.

Interestingly, evidence for ungulates suppressing shrub growth was only observed at intermediate summer temperatures, and even at these temperatures the effect was minor. Forage preferences and landscape use of the ungulates may explain these results, as well as favorable climatic conditions that may enable effective compensatory growth of shrubs. Earlier local studies have shown that ungulates may counteract the impacts of warming on tundra shrub growth, but we demonstrated that ungulates’ potential to suppress shrub growth is not always evident and may be limited to certain temperature and precipitation conditions.

Reference: Vuorinen, K., Austrheim, G., Tremblay, J.-P., Myers-Smith, I.H., Hortman, H.I., Frank, P., Barrio, I.C., Dalerum, F., Björkman, M.P., Björk, R.G., Ehrich, D., Sokolov, A., Sokolova, N., Ropars, P., Boudreau, S., Normand, S., Prendin, A.L., Schmidt, N.M., Pacheco, A., Post, E., John, C., Kerby, J.T., Sullivan, P.F., Le Moullec, M., Hansen, B.B., Van der Wal, R., Pedersen, Å.Ø., Sandal, L., Gough, L., Young, A., Li, B., Magnússon, R.Í., Sass-Klaassen, U., Buchwal, A., Welker, J.M., Grogan, P., Andruko, R., Morrissette-Boileau, C., Volkovitskiy, A., Terekhina, A., Speed, J.D.M., 2022. Growth rings show limited evidence for ungulates’ potential to suppress shrubs across the Arctic. Environmental Research Letters. https://iopscience.iop.org/article/10.1088/1748-9326/ac5207/meta


This paper is a contribution to the DISENTANGLE project and was included as one of the chapters of Katariina’s PhD at the Norwegian University of Science and Technology.

Photograph: Muskoxen (Ovibos moschatus) in a willow patch (photo: Katariina Vuorinen)

A protocol for conducting a systematic review on the effects of herbivore diversity on tundra ecosystems

How do the effects of different herbivores combine to affect tundra ecosystem? This is the question we want to address with a new systematic review. This project builds on earlier work by Herbivory Network members where we compiled available evidence on effects of herbivores on tundra vegetation using a systematic map (Soininen et al, 2021). The systematic review will expand this effort beyond tundra vegetation to include other ecosystem components as well, and will focus on studies that compare areas with different herbivore diversity.

Following the guidelines of the Collaboration for Environmental Evidence (CEE), the process of the systematic review involves two steps: developing a systematic review protocol (that is then peer-reviewed and published), and using that protocol to conduct the actual systematic review. A smaller group of authors has been developing the protocol, that has now been published in Environmental Evidence. We have also started to work on the systematic review. There is still time if you want to get involved in this project! See here the call for collaboration, and contact Laura Barbero-Palacios for more information.

You can access the full text here.

Reference: Barrio, I.C., Barbero-Palacios, L., Kaarlejärvi, E., Speed, J.D.M., Heiðmarsson, S., Hik, D.S., Soininen, E.M. (2022) What are the effects of herbivore diversity on tundra ecosystems? A systematic review protocol. Environmental Evidence 11:1. https://doi.org/10.1186/s13750-022-00257-z

This project is a contribution to the TUNDRAsalad and CHARTER projects.

Will borealization of Arctic tundra herbivore communities be driven by climate warming or vegetation change?

Arctic ecosystems are changing fast, and these changes involve both plant and herbivore communities. Poleward shifts of species distributions, termed borealization when referring to northward movements of boreal species into tundra, have been observed and are expected to continue. Increasing greenness, shrubification and treeline advance in Arctic ecosystems have been associated with warming trends. Vertebrate herbivores have the potential to limit greening and shrub advance and expansion on the tundra, posing the question of whether changes in herbivore communities could partly mediate the impacts of climate warming on Arctic tundra. Therefore, future changes in the herbivore community in the Arctic tundra will depend on whether the community tracks the changing climates directly (i.e. occurs in response to temperature) or indirectly, in response to vegetation changes (which can be modified by trophic interactions).

We used biogeographic and remotely sensed data to quantify spatial variation in vertebrate herbivore communities across the boreal forest and Arctic tundra biomes and assess whether borealization of vertebrate herbivore communities is a direct response to warming temperatures, or an indirect response through changing vegetation. We then tested whether present-day herbivore community structure is determined primarily by temperature or vegetation.

The composition of herbivore communities across the biome boundary between the boreal forest and the tundra is mainly driven by temperature, rather than vegetation productivity and woody plant cover.

We found that vertebrate herbivore communities are significantly more diverse in the boreal forest than in the Arctic tundra in terms of species richness, phylogenetic diversity and functional diversity. A clear shift in community structure was observed at the biome boundary, with stronger northward declines in diversity in the Arctic tundra. Interestingly, important functional traits characterizing the role of herbivores in limiting tundra vegetation change, such as body mass and woody plant feeding, did not show threshold changes across the biome boundary. The composition of herbivore communities was mainly driven by temperature rather than by vegetation productivity or woody plant cover. Thus, our study does not support the premise that herbivore-driven limitation of Arctic tundra shrubification or greening would limit herbivore community change in the tundra. Instead, borealization of tundra herbivore communities is likely to result from the direct effect of climate warming.

You can access the full text here.

Reference: Speed, J.D., Chimal‐Ballesteros, J.A., Martin, M.D., Barrio, I.C., Vuorinen, K.E. and Soininen, E.M., 2021. Will borealization of Arctic tundra herbivore communities be driven by climate warming or vegetation change?. Global Change Biology27(24):6568-6577.


Photo: reindeer grazing (photo by Hannes Skarin)

Silicon-based defence and nutrient levels in tundra grasses under herbivory

To what extent herbivores concomitantly alter silicon-based defence and nutrient levels in tundra grasses will ultimately determine changes in the quality of their pastures. In this study, we asked to what extent keystone tundra herbivores, small rodents and reindeer, affect silicon content and silicon:nutrient ratios of grasses found in tundra-grasslands.

Herbivores did not promote a net silicon accumulation in grasses, but rather enhanced their quality by increasing leaf nitrogen and phosphorus levels, thus decreasing silicon:nutrient ratios. Yet, the magnitude of these quality increments varied depending on the herbivore(s) involved and differed between inherently silicon-rich and silicon-poor grasses, ultimately leading to the formation of a fine-scale mosaic of tundra-patches with different nutrient values. In tundra-patches utilised by both herbivores, the quality of inherently silicon-rich grasses was further decreased relative to that of the already more palatable silicon-poor grasses. This could provide an advantage against herbivory, potentially being one of the pathways through which tundra-grassland vegetation states dominated by silicon-rich grasses are generally maintained by herbivores.

Reference: Petit Bon, M., Inga, K.G., Utsi, T.A., Jónsdóttir, I.S. and Bråthen, K.A. (2022), Forage quality in tundra grasslands under herbivory: Silicon-based defences, nutrients, and their ratios in grasses. Journal of Ecology 110(1): 129-143. https://doi.org/10.1111/1365-2745.13790


Text: Matteo Petit Bon, Czech Academy of Sciences.

Picture: The fence built in the 1950s at Ifjordfjellet, Finnmark, separates spring/fall migratory and summer pasture ranges for semi-domesticated reindeer (Photo: Matteo Petit-Bon).

Vegetation and soil C/N chemistry is differentially affected by perturbations like goose grubbing and warming in the high-Arctic

As vegetation and soil jointly regulate whole-ecosystem processes, differential sensitivity and magnitude of their carbon and nitrogen responses to environmental perturbations may have implications for the functioning of tundra ecosystems. In this study, we addressed sensitivity and magnitude of short-term carbon and nitrogen responses of vascular plants, mosses, and soil to simulated goose disturbance and warming across three habitats that differ in soil moisture in a high-Arctic ecosystem in Svalbard.

Though the system’s total carbon and nitrogen contents were relatively resistant to both perturbations, the three studied ecosystem compartments still differed in their chemical responses (vascular plants > soil > mosses), and such differential sensitivity was further exacerbated by their different responses across habitats (mesic > moist > wet).

These findings highlight the potential for environmental perturbations to have small, yet differential short-term impacts on the carbon and nitrogen contents of vascular plants, mosses, and soil, both within and between tundra-habitats. They also imply that assessments of a single ecosystem compartment in a given context cannot be extrapolated to the whole ecosystem, suggesting that addressing vegetation and soil chemical responses in different habitats can advance our predictive capability of how the biogeochemistry of tundra ecosystems respond to environmental changes.

Reference: Petit Bon, M., Böhner, H., Bråthen, K. A., Ravolainen, V. T., and Jónsdóttir, I. S.. 2021. Variable responses of carbon and nitrogen contents in vegetation and soil to herbivory and warming in high-Arctic tundra. Ecosphere 12( 9):e03746. https://doi.org/10.1002/ecs2.3746


Text: Matteo Petit Bon, Czech Academy of Sciences.

Picture: Pink-footed geese after their arrival in Svalbard (Photo: Matteo Petit-Bon).

Synergistic effects of insect herbivory and changing climate on plant volatile emissions in the subarctic tundra

Insect herbivory is a significant source of plant stress and, due to climate change, the herbivory stress is continuously increasing. In the Arctic, where climate warming proceeds with the rate twice of the global average, insect herbivory stress to vegetation is expected to become more severe. In response to herbivory, plants produce specialized metabolites to fulfil physiological and ecological functions, which include volatile organic compounds (VOCs). VOCs are highly reactive and play an important role in biosphere-atmosphere interactions. Various biotic and abiotic stressors, including insect herbivory, can enhance and/or induce the production of many VOCs and alter the composition of the overall VOC blend emitted from the plants.

Due to lack of the empirical studies in the Arctic, it is unclear how the effects of insect herbivory and changing climate such as warming and increased cloudiness, which are both predicted conditions for the future Arctic, will affect VOC emissions in the changing Arctic.

Therefore, in this study we assessed how experimental manipulations of temperature and light availability in subarctic tundra, that had been maintained for 30 years at the time of the measurements, affect the VOC emissions from a widespread dwarf birch when subjected to herbivory by local geometrid moth larvae, the autumnal moth and the winter moth.

Our study showed that whether the future subarctic tundra in Northern Fennoscandia experiences rising temperatures or increasing cloudiness, and therefore milder temperature increases, matter very little for herbivory-related VOC emissions. Under both conditions, we showed strong interactions with herbivory effects on VOC emissions with potentially positive feedbacks on cloud formation. We also show that acclimation of plants to long-term climate treatments, which has resulted in changes in anatomical traits, might strongly interact with volatile responses to insect herbivory. This finding further complicates predictions of how climate change, together with interacting biotic stresses, affects VOC emissions in the Arctic.

Full article here:  https://doi.org/10.1111/gcb.15773

Reference: Rieksta, J., Li, T., Michelsen, A., & Rinnan, R. (2021). Synergistic effects of insect herbivory and changing climate on plant volatile emissions in the subarctic tundra. Global Change Biology, 27(20), 5030-5042.


Text: Jolanta Rieksta, University of Copenhagen.

Picture: Measuring volatile organic compound (VOC) emissions from dwarf birch in the field using branch enclosure method where VOC are collected using custom built pull-push systems. (Photo: Jolanta Rieksta).

The paradox of forbs in grasslands and the legacy of the mammoth steppe

Have you ever been puzzled by the fact that we as ecologists often lump a range of plant species into a single functional category? In this paper the focus is to the most species rich functional grouping we often apply in grasslands, the herbaceous plants with colorful flowers, the forbs. We lump them because each single species often make up a very small abundance, yet why are there so many forb species with small abundances? And why are they in sum often inferior in abundance to the grasses? If looking at forbs in a paleoecological perspective, in the cold Mammoth steppe and in which DNA-based evidence indicates the forbs flourished, we may rethink the importance of forbs and why they often do not seem to thrive in grasslands today.

Hypothesized plant–megafauna feedbacks in the mammoth steppe

You can access the paper here: https://esajournals.onlinelibrary.wiley.com/doi/10.1002/fee.2405

Reference: Bråthen, K.A., Pugnaire, F.I. and Bardgett, R.D., 2021. The paradox of forbs in grasslands and the legacy of the mammoth steppe. Frontiers in Ecology and the Environment.


Text: Kari Anne Bråthen, Professor, UiT The Arctic University of Norway.
Illustration: Ernst Asbjørn Høgtun

Systematic map of herbivore studies in the Arctic

Herbivores modify arctic vegetation and can counteract some of the climate-change driven increases in shrub cover and treeline advance induced by climate-change in the tundra. However, the effects of herbivores on plants and ecosystem structure and function vary across the Arctic and seem to depend on the environmental conditions under which herbivory takes place. Therefore, studies in one location can lead to different results than studies in another location, and generalizing the impacts of herbivores becomes difficult. Our systematic map assessed how well the existing literature of herbivore impacts on vegetation covers the environmental variation in the Arctic, to understand how robust are the conclusions that we can make about the effects of herbivores across the tundra biome. Our results show that herbivory research is concentrated in parts of the Arctic that are warmer, wetter, near the coast and that have experienced a moderate increase in temperature. The current evidence base might thus provide an incomplete picture of the effects of herbivores on Arctic vegetation throughout the region.

The database of studies of herbivore effects on arctic vegetation is available through an interactive visualization tool (Arctic Herbivory Systematic Map) that allows exploration of individual environmental variables and the coded data.

This systematic map has been a large effort led by Eeva Soininen. You can find the paper here.

Reference: Soininen, E.M., Barrio, I.C., Bjørkås, R., Björnsdóttir, K., Ehrich, D., Hopping, K.A., Kaarlejärvi, E., Kolstad, A.L., Abdulmanova, S., Björk, R.G.,, Bueno, C.G., Eischeid, I.,, Finger-Higgens, R., Forbey, J.S., Gignac, C., Gilg, O.,, den Herder, M., Holm, H.S., Hwang, B.C., Jepsen, J.U., Kamenova, S.,, Kater, I., Koltz, A.M.,, Kristensen, J.A.,, Little, C.J., Macek, P.,, Mathisen, K.M., Metcalfe, D.B.,, Mosbacher, J.B., Mörsdorf, M., Park, T.,, Propster, J.R.,, Roberts, A.J., Serrano, E., Spiegel, M.P., Tamayo, M., Tuomi, M.W., Verma, M., Vuorinen, K.E.M., Väisänen, M., van der Wal, R., Wilcots, M.E., Yoccoz, N.G., Speed, J. D. (2021). Location of studies and evidence of effects of herbivory on Arctic vegetation: a systematic map. Environmental Evidence, 10:25. https://doi.org/10.1186/s13750-021-00240-0