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Global vegetation regimes vary in belowground carbon (C) and nitrogen (N) dynamics. However, disentangling large-scale climatic controls from the effects of intrinsic plant–soil–microbial feedbacks on belowground processes is challenging. In local gradients with similar pedo-climatic conditions, effects of plant–microbial feedbacks may be isolated from large-scale drivers. Across a subarctic–alpine mosaic of historic grazing fields and surrounding heath and birch forest, we evaluated whether vegetation-specific plant–microbial feedbacks involved contrasting N cycling characteristics and C and N stocks in the organic topsoil. We sequenced soil fungi, quantified functional genes within the inorganic N cycle, and measured 15N natural abundance. In grassland soils, large N stocks and low C : N ratios associated with fungal saprotrophs, archaeal ammonia oxidizers, and bacteria capable of respiratory ammonification, indicating maintained inorganic N cycling a century after abandoned reindeer grazing. Toward forest and heath, increasing abundance of mycorrhizal fungi co-occurred with transition to organic N cycling. However, ectomycorrhizal fungal decomposers correlated with small soil N and C stocks in forest, while root-associated ascomycetes associated with small N but large C stocks in heath, uncoupling C and N storage across vegetation types. We propose that contrasting, positive plant–microbial feedbacks stabilize vegetation trajectories, resulting in diverging soil C : N ratios at the landscape scale. publishedVersion

Open Science (OS) comprises a variety of practices and principles that are broadly intended to improve the quality and transparency of research, and the concept is gaining traction. Since OS has multiple facets and still lacks a unifying definition, it may be interpreted quite differently among practitioners. Moreover, successfully implementing OS broadly throughout science requires a better understanding of the conditions that facilitate or hinder OS engagement, and in particular, how practitioners learn OS in the first place. We addressed these issues by surveying OS practitioners that attended a workshop hosted by the Living Norway Ecological Data Network in 2020. The survey contained scaled-response and open-ended questions, allowing for a mixed-methods approach. Out of 128 registered participants we obtained survey responses from 60 individuals. Responses indicated usage and sharing of data and code, as well as open access publications, as the OS aspects most frequently engaged with. Men and those affiliated with academic institutions reported more frequent engagement with OS than women and those with other affiliations. When it came to learning OS practices, only a minority of respondents reported having encountered OS in their own formal education. Consistent with this, a majority of respondents viewed OS as less important in their teaching than in their research and supervision. Even so, many of the respondents’ suggestions for what would help or hinder individual OS engagement included more knowledge, guidelines, resource availability and social and structural support; indicating that formal instruction can facilitate individual OS engagement. We suggest that the time is ripe to incorporate OS in teaching and learning, as this can yield substantial benefits to OS practitioners, student learning, and ultimately, the objectives advanced by the OS movement.

1. The potential of large mammalian herbivores to shift plant communities between nitrogen (N) and phosphorus (P) limitation has received little attention so far. However, herbivores can influence the cycling of these growth‐limiting nutrients, and thereby affect plant nutrient limitation and productivity. Tundra ecosystems are nutrient‐poor and commonly grazed by large herbivores like reindeer and may thus be responsive to such changes. 2. Here, we examined the effect of long‐term light and heavy reindeer grazing on nutrient limitation of plant growth in a Scandinavian arctic tundra. We are the first to conduct a factorial N and P fertilization experiment across the two grazing regimes in two functionally contrasting vegetation types: heath and meadow. 3. Annual primary productivity (APP) showed contrasting responses to our fertilization treatments under light and heavy grazing. Under light grazing, APP increased in response to N + P additions in both the heath and meadow. Under heavy grazing, APP increased in response to N in the heath, with an additional positive effect of N + P combined, while APP increased in response to P and N + P additions in the meadow. 4. These results clearly show that an increase in the grazing intensity of reindeer facilitated a shift towards more P‐limited conditions in Scandinavian arctic tundra, by increasing N cycling without having a corresponding positive effect on P cycling. In the N‐poor heath, reindeer increased soil N availability at least partly due to a shift towards more N‐rich graminoids, while in the meadow, reindeer decreased soil P availability. The mechanisms behind this decrease remain unclear, but reindeer may simply export more P from the system than N due to their large P demand for the production of their antlers. 5. Synthesis. We conclude that heavy and long‐term reindeer grazing promoted a more P‐limited tundra, thus experimentally confirming the potential of large mammalian herbivores to influence nutrient limitation of plant growth.

Question Herbivores exert strong influences on vegetation through activities such as trampling, defoliation, and fertilization. The combined effect of these activities on plant performance may cause dramatic vegetation shifts. Because herbivore pressures and the relative importance of their different activities are not equally distributed across the landscape, it is important to understand their isolated effect. One example of an herbivore-induced vegetation shift is the reindeer-driven transition from a subarctic tundra vegetation dominated by dwarf shrubs into a more productive, graminoid-dominated state. Here, we asked how each of the grazing activities by reindeer separately and combined shape vegetation composition. Location Nordreisa, Norway. Methods We used a field experiment over six summers to study the separate and interacting effects of reindeer trampling, defoliation, addition of faeces and removal of moss on tundra heath vegetation, and to identify which of these factors were most important in driving the plant community towards a graminoid-dominated state. Results The combination of all treatments resulted in the strongest changes in vegetation, but trampling was the single most important factor altering the vegetation composition by reducing the abundance of both evergreen and deciduous dwarf shrubs. In contrast to what was expected, none of our treatments, separate or combined, resulted in an increased abundance of graminoids in 5 years, although such rapid vegetation changes have been observed in the field in similar environmental conditions. Conclusions Trampling is the key process by which reindeer influence the abundance of functional groups, but only many processes combined result in strong changes in community composition. Moreover, additional factors not included in this experiment, such as urine, may be important in causing a state shift to a graminoid-dominated community. publishedVersion

Invertebrate herbivores depend on external temperature for growth and metabolism. Continued warming in tundra ecosystems is proposed to result in increased invertebrate herbivory. However, empirical data about how current levels of invertebrate herbivory vary across the Arctic is limited and generally restricted to a single host plant or a small group of species, so predicting future change remains challenging. We investigated large-scale patterns of invertebrate herbivory across the tundra biome at the community level and explored how these patterns are related to long-term climatic conditions and year-of-sampling weather, habitat characteristics, and aboveground biomass production. Utilizing a standardized protocol, we collected samples from 92 plots nested within 20 tundra sites during summer 2015. We estimated the community-weighted biomass lost based on the total leaf area consumed by invertebrates for the most common plant species within each plot. Overall, invertebrate herbivory was prevalent at low intensities across the tundra, with estimates averaging 0.94% and ranging between 0.02 and 5.69% of plant biomass. Our results suggest that mid-summer temperature influences the intensity of invertebrate herbivory at the community level, consistent with the hypothesis that climate warming should increase plant losses to invertebrates in the tundra. However, most of the observed variation in herbivory was associated with other site level characteristics, indicating that other local ecological factors also play an important role. More details about the local drivers of invertebrate herbivory are necessary to predict the consequences for rapidly changing tundra ecosystems. - 2019, Springer-Verlag GmbH Germany, part of Springer Nature. This study is a contribution of the Herbivory Network ( https://herbivory.lbhi.is ). SIR and DSH were supported by the Natural Sciences and Engineering Research Council of Canada (RGPNS-305471); ICB was supported by a postdoctoral fellowship funded by the Icelandic Research Fund (Rannsóknasjóður, Grant nr 152468–051) and AXA Research Fund (15-AXA-PDOC-307); JMA was supported by Carl Tryggers Stiftelse för Vetenskaplig Forskning (Sweden) and Qatar Petroleum; MVK and VZ were supported by the Academy of Finland (Project 276671); AS and NS were supported by the Grant of Russian fund for basic research (18-05-60261); Field work on Bylot Island was supported by the Natural Sciences and Engineering Research Council of Canada, the network of Center of excellence ArcticNet, and the Polar Continental Shelf Program to GG and EL; SK and the Burntpoint Creek Research Station was supported by the Waterfowl, Wetlands and Far North research unit of the Ontario Ministry of Natural Resources and Forestry, with assistance from K. Bennet, R. Brook, and L. Pollock; O. Krasheninnikova assisted with the collection plant samples on Erkuta; SS was partly supported by a state assignment of the Institute of Plant and Animal Ecology UB RAS; DDE was supported by a Grant to JO from the Swedish Research Council Formas 2012–1039, 2012–230, 2015–1091 and a Grant to DDE from Gunnar and Ruth Björkmans fund for botanical research in northern Sweden; Sampling in Svalbard was conducted during an UNIS field course (ISJ); JSP was supported by the WSL Institute for Snow and Avalanche Research, and an INTERACT Transnational Access Research Grant; Research at Toolik Field Station by ALA was supported by National Science Foundation Grants DEB 102683 to support the Arctic LTER and OPP 0908502; JDMS was supported by the Research Council of Norway (262064); C. Brachmann assisted with Fig. 1 ; H.C. Proctor and B.A. Keddie assisted with gall damage determination. Scopus

Ecosystems where severe disturbance has induced permanent shifts in vegetation and soil processes may represent alternative stable states. To date, little is known on how long-lasting changes in soil processes are following such disturbances, and how the changes in plant and soil processes between the alternative states eventually manifest themselves in soil organic matter (SOM) storage. Here, we analyzed plant density, the shrub : forb ratio, microbial respiration, extracellular enzyme activities and SOM stocks in soils of subarctic tundra and historical milking grounds, where reindeer herding induced a vegetation transition from deciduous shrubs to graminoids several centuries earlier but were abandoned a century ago. This provides the possibility to compare sites with similar topography, but highly contrasting vegetation for centuries. We found that enzymatic activities and N:P stoichiometry differed between control and disturbed sites, confirming that culturally induced vegetation shifts exert lasting impacts on tundra soil processes. Transition zones, where shrubs had encroached into the historical milking grounds during the past 50 yr, indicated that microbial activities for N and P acquisition changed more rapidly along a vegetation shift than those for microbial C acquisition. Although plant and soil processes differed between control and disturbed sites, we found no effect of historical vegetation transition on SOM stock. Across the study sites, soil SOM stocks were correlated with total plant density but not with the shrub : forb ratio. Our finding that SOM stock was insensitive to a centennial difference in plant community composition suggests that, as such, grazing-induced alternative vegetation states might not necessarily differ in SOM sequestration.

The above mentioned article was originally scheduled for publication in the special issue on Ecology of Tundra Arthropods with guest editors Toke T. Høye . Lauren E. Culler. Erroneously, the article was published in Polar Biology, Volume 40, Issue 11, November, 2017. The publisher sincerely apologizes to the guest editors and the authors for the inconvenience caused.

Historical contingency is the impact of past events, like the timing and order of species arrival, on community assembly, and can sometimes result in alternative stable states of ecological communities. Large herbivores, wild and domestic, can cause profound changes in the structure and functioning of plant communities and therefore probably influence historical contingency; however, little empirical data on the stability of such shifts or subsequent drivers of stability are available. We studied the centennial legacy of reindeer (Rangifer tarandus) pressure on arctic tundra vegetation by considering historical milking grounds (HMGs): graminoid- and forb-dominated patches amid shrub-dominated tundra, formed by historical Sami reindeer herding practices that ended approximately 100 years ago. Our results show that the core areas of all studied HMGs remained strikingly stable, being hardly invaded by surrounding shrubs. Soil nitrogen concentrations were comparable to heavily grazed areas. However, the HMGs are slowly being reinvaded by vegetative growth of shrubs at the edges, and the rate of ingrowth increased with higher mineral N availability. Furthermore, our data indicate that several biotic feedbacks contribute to the stability of the HMGs: increased nutrient turnover supporting herbaceous vegetation, strong interspecific competition preventing invasion and herbivore damage to invading shrubs. In particular, voles and lemmings appear to be important, selectively damaging shrubs in the HMGs. We concluded that HMGs provide clear evidence for historical contingency of herbivore effects in arctic ecosystems. We showed that several biotic feedbacks can contribute to subsequent vegetation stability, but their relative importance will vary in time and space. Originnally included in thesis in manuscript form

Chronic, low intensity herbivory by invertebrates, termed background herbivory, has been understudied in tundra, yet its impacts are likely to increase in a warmer Arctic. The magnitude of these changes is however hard to predict as we know little about the drivers of current levels of invertebrate herbivory in tundra. We assessed the intensity of invertebrate herbivory on a common tundra plant, the dwarf birch (Betula glandulosa-nana complex), and investigated its relationship to latitude and climate across the tundra biome. Leaf damage by defoliating, mining and gall-forming invertebrates was measured in samples collected from 192 sites at 56 locations. Our results indicate that invertebrate herbivory is nearly ubiquitous across the tundra biome but occurs at low intensity. On average, invertebrates damaged 11.2% of the leaves and removed 1.4% of total leaf area. The damage was mainly caused by external leaf feeders, and most damaged leaves were only slightly affected (12% leaf area lost). Foliar damage was consistently positively correlated with mid-summer (July) temperature and, to a lesser extent, precipitation in the year of data collection, irrespective of latitude. Our models predict that, on average, foliar losses to invertebrates on dwarf birch are likely to increase by 6–7% over the current levels with a 1 °C increase in summer temperatures. Our results show that invertebrate herbivory on dwarf birch is small in magnitude but given its prevalence and dependence on climatic variables, background invertebrate herbivory should be included in predictions of climate change impacts on tundra ecosystems. This study is a joint contribution of the Herbivory Network (http://herbivory.biology.ualberta.ca) and the Network for Arthropods of the Tundra (NeAT; https://tundraarthropods.wordpress.com/). Dwarf birch distribution maps were kindly provided by Kyle Joly. Sample collection during 2014 was facilitated by INTERACT (http://www.eu-interact.org/). ICB was supported by a postdoctoral fellowship funded by the Icelandic Research Fund (Rannsóknasjóður, grant nr 152468-051) and AXA Research Fund (15-AXA-PDOC-307); MtB and EK were supported by the Nordic Centre of Excellence TUNDRA, funded by the Norden Top-Level Research Initiative ‘‘Effect Studies and Adaptation to Climate Change’’; EMS and KAB were supported by COAT (Climate-ecological Observatory of the Arctic Tundra); AB was supported by MOBILITY PLUS (1072/MOB/2013/0) and the Polish-American Fulbright Commission; CGB was supported by IUT 20-28, EcolChang e Center of Excellence; BCF and TK were supported by the Academy of Finland (project 256991); MMPDH was supported by The Netherlands Organization for Scientific Research (NWO-ALW, VIDI grant 864.09.014); DSH was supported by the Natural Sciences and Engineering Research Council of Canada; AH was supported by the Research Council of Norway (grant 244557/E50); JL was funded by the German Research Foundation DFG (project WI 2680/8-1); MM-F was supported by a NERC IRF fellowship NE/L011859/1; SN was supported by the Villum foundation’s Young Investigator Programme (VKR023456); JS was supported by Kempestiftelserna and the Research Foundation Flanders (FWO); AS and NS were supported by the grant of RFBR (project 16-44-890108), grant of UB of RAS (project 15-15-4-35) and IEC “Arctic” of Yamal Government Department of Science and Innovation; LES and PAW were supported by the UK Natural Environment Research Council (NERC) grant NE/K000284/1; MVK and VZ were supported by the Academy of Finland (project 276671). Scopus

How can we best convey the principles of open science to students, make use of open science in teaching and mentoring, and develop the tools and techniques for the future of open science? This workshop is designed to share and collate experiences and ideas in the intersection of open science and education, and forecast the way forward. The workshop is a collaboration between Living Norway andbioCEED, located at the University of Bergen. The presentation was the introduction to the workshop