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Foundation species provide habitat and modify the availability of resources to other species. In nature, multiple foundation species may occur in mixture, but little is known on how their interactions shape the community assembly of associated species. Lichens provide both structural habitat and resources to a variety of associated organisms and thereby serve as foundation species. In this study, we use mat-forming lichens and their associated micro-arthropods as a miniature ecosystem to study potential synergies between foundation species diversity and the abundance and functional diversity of higher trophic levels. We created lichen patches with monocultures and mixtures of up to four species, and extracted Collembola (identified to species level), Oribatida, Mesostigmata, Pseudoscorpiones, and Araneae with Tullgren apparatuses after 106 days of incubation within a natural lichen mat. We found that different lichen species supported different arthropod abundances. For 19 out of a total of 55 lichen mixtures and arthropod groups, we found non-additive, synergistic effects on arthropod abundance, although the specific lichen mixture causing synergistic effects differed with arthropod group. In addition, synergistic effects on arthropod abundance were more common for arthropod groups at lower trophic levels. The functional diversity of lichen mixtures explained patterns in Collembola abundance, but in the opposite direction than hypothesized because synergistic responses were more frequent in functionally similar lichen mixtures. Finally, we found few effects of lichen mixture identity or diversity on the functional diversity of Collembola communities. When applied to large-scale ecosystems, our results suggest that understanding interactions between coexisting foundation species and identifying those species that drive synergistic effects of foundation species on consumer biota, is likely to be of importance to biodiversity conservation and restoration efforts. Published version This study was funded by a grant from the Norwegian Research Counsel (249,902/F20) to JA.

Data and code in support of our paper "Comparing Global Sentinel-2 Land Cover Maps for Regional Species Distribution Modeling" published in Remote Sensing. The 'Venter et al R script.R' can be used to reproduce the analysis. It will reference the CSV and GeoTIFF files hosted in this repository. The abstract from the paper: Mapping the spatial and temporal dynamics of species distributions is necessary for biodiversity conservation land-use planning decisions. Recent advances in remote sensing and machine learning have allowed for high-resolution species distribution modeling that can inform landscape-level decision-making. Here we compare the performance of three popular Sentinel-2 (10-m) land cover maps, including dynamic world (DW), European land cover (ELC10), and world cover (WC), in predicting wild bee species richness over southern Norway. The proportion of grassland habitat within 250 m (derived from the land cover maps), along with temperature and distance to sandy soils, were used as predictors in both Bayesian regularized neural network and random forest models. Models using grassland habitat from DW performed best (RMSE = 2.8 ± 0.03; average ± standard deviation across models), followed by ELC10 (RMSE = 2.85 ± 0.03) and WC (RMSE = 2.87 ± 0.02). All satellite-derived maps outperformed a manually mapped Norwegian land cover dataset called AR5 (RMSE = 3.02 ± 0.02). When validating the model predictions of bee species richness against citizen science data on solitary bee occurrences using generalized linear models, we found that ELC10 performed best (AIC = 2278 ± 4), followed by WC (AIC = 2367 ± 3), and DW (AIC = 2376 ± 3). While the differences in RMSE we observed between models were small, they may be significant when such models are used to prioritize grassland patches within a landscape for conservation subsidies or management policies. Partial dependencies in our models showed that increasing the proportion of grassland habitat is positively associated with wild bee species richness, thereby justifying bee conservation schemes that aim to enhance semi-natural grassland habitat. Our results confirm the utility of satellite-derived land cover maps in supporting high-resolution species distribution modeling and suggest there is scope to monitor changes in species distributions over time given the dense time series provided by products such as DW.

In the high Arctic, plant community species composition generally responds slowly to climate warming, whereas less is known about the community functional trait responses and consequences for ecosystem functioning. Slow species turnover and large distribution ranges of many Arctic plant species suggest a significant role of intraspecific trait variability in functional responses to climate change. Here, we compare taxonomic and functional community compositional responses to a long-term (17 years) warming experiment in Svalbard, replicated across three major high Arctic habitats shaped by topography and contrasting snow regimes. We observed taxonomic compositional changes in all plant communities over time. Still, responses to experimental warming were minor and most pronounced in the drier habitats with relatively early snowmelt timing and long growing seasons (Cassiope and Dryas heaths). The habitats were clearly separated in functional trait space, defined by twelve size- and leaf economics-related traits, primarily due to interspecific trait variation. Functional traits also responded to experimental warming, most prominently in the Dryas heath and mostly due to intraspecific trait variation. Leaf area and leaf mass increased, and leaf δ15N decreased in response to the warming treatment. Intraspecific trait variability ranged between 30% and 71% of the total trait variation, reflecting functional resilience of those communities, dominated by long-lived plants, due to either phenotypic plasticity or genotypic variation that most likely underlies the observed resistance of high Arctic vegetation to climate warming. We further explored the consequences of trait variability for ecosystem functioning by measuring peak season CO2 fluxes. Together, environmental, taxonomic, and functional trait variables explained a large proportion of the variation in net ecosystem exchange (NEE), which increased when intraspecific trait variation was accounted for. In contrast, even though ecosystem respiration and gross ecosystem production both increased in response to warming across habitats, they were mainly driven by the direct kinetic impacts of temperature on plant physiology and biochemical processes. Our study shows that long-term experimental warming has a modest but significant effect on plant community functional trait composition and suggests that intraspecific trait variability is a key feature underlying high Arctic ecosystem resistance to climate warming.

The International Tundra Experiment (ITEX) was established to understand how environmental change impacts Arctic and alpine ecosystems. The success of the ITEX-network has allowed for several important across-site syntheses, and for some ITEX-sites enough data have now been collected to perform within-site syntheses on the effects of environmental change across ecological scales. In this study, we analyze climate data and synthesize three decades of research on the ecological effects of environmental change at the ITEX-site at Finse, southern Norway. We found a modest warming rate of +0.36 °C per decade and minor effects on growing season length. Maximum winter snow depth was highest in winters with a positive North Atlantic Oscillation. Our synthesis included 80 ecological studies from Finse, biased towards primary producers with few studies on ecological processes. Species distributions depended on microtopography and microclimate. Experimental warming had contrasting effects on abundance and traits of individual species and only modest effects at the community-level above and below ground. In contrast, nutrient addition experiments caused strong responses in primary producer and arthropod communities. This within-site synthesis enabled us to conclude how different environmental changes (experimental and ambient warming, nutrient addition, and environmental gradients) impact across ecological scales, which is challenging to achieve with across-site approaches.

Mapping the spatial and temporal dynamics of species distributions is necessary for biodiversity conservation land-use planning decisions. Recent advances in remote sensing and machine learning have allowed for high-resolution species distribution modeling that can inform landscape-level decision-making. Here we compare the performance of three popular Sentinel-2 (10-m) land cover maps, including dynamic world (DW), European land cover (ELC10), and world cover (WC), in predicting wild bee species richness over southern Norway. The proportion of grassland habitat within 250 m (derived from the land cover maps), along with temperature and distance to sandy soils, were used as predictors in both Bayesian regularized neural network and random forest models. Models using grassland habitat from DW performed best (RMSE = 2.8 ± 0.03; average ± standard deviation across models), followed by ELC10 (RMSE = 2.85 ± 0.03) and WC (RMSE = 2.87 ± 0.02). All satellite-derived maps outperformed a manually mapped Norwegian land cover dataset called AR5 (RMSE = 3.02 ± 0.02). When validating the model predictions of bee species richness against citizen science data on solitary bee occurrences using generalized linear models, we found that ELC10 performed best (AIC = 2278 ± 4), followed by WC (AIC = 2367 ± 3), and DW (AIC = 2376 ± 3). While the differences in RMSE we observed between models were small, they may be significant when such models are used to prioritize grassland patches within a landscape for conservation subsidies or management policies. Partial dependencies in our models showed that increasing the proportion of grassland habitat is positively associated with wild bee species richness, thereby justifying bee conservation schemes that aim to enhance semi-natural grassland habitat. Our results confirm the utility of satellite-derived land cover maps in supporting high-resolution species distribution modeling and suggest there is scope to monitor changes in species distributions over time given the dense time series provided by products such as DW. pollinators; grassland; wild bees; management; conservation; spatial modeling

This article comments on:Max Mallen-Cooper, Bente J. Graae and Will K. Cornwell. Lichens buffer tundra microclimate more than the expanding shrub Betula nana, Annals of Botany, Volume 128, Issue 4, 3 September 2021, Pages 407–418, https://doi.org/10.1093/aob/mcab041

Publisher's version (útgefin grein), Climatic impacts are especially pronounced in the Arctic, which as a region is warming twice as fast as the rest of the globe. Here, we investigate how mean climatic conditions and rates of climatic change impact parasitoid insect communities in 16 localities across the Arctic. We focus on parasitoids in a widespread habitat, Dryas heathlands, and describe parasitoid community composition in terms of larval host use (i.e., parasitoid use of herbivorous Lepidoptera vs. pollinating Diptera) and functional groups differing in their closeness of host associations (koinobionts vs. idiobionts). Of the latter, we expect idiobionts—as being less fine‐tuned to host development—to be generally less tolerant to cold temperatures, since they are confined to attacking hosts pupating and overwintering in relatively exposed locations. To further test our findings, we assess whether similar climatic variables are associated with host abundances in a 22 year time series from Northeast Greenland. We find sites which have experienced a temperature rise in summer while retaining cold winters to be dominated by parasitoids of Lepidoptera, with the reverse being true for the parasitoids of Diptera. The rate of summer temperature rise is further associated with higher levels of herbivory, suggesting higher availability of lepidopteran hosts and changes in ecosystem functioning. We also detect a matching signal over time, as higher summer temperatures, coupled with cold early winter soils, are related to high herbivory by lepidopteran larvae, and to declines in the abundance of dipteran pollinators. Collectively, our results suggest that in parts of the warming Arctic, Dryas is being simultaneously exposed to increased herbivory and reduced pollination. Our findings point to potential drastic and rapid consequences of climate change on multitrophic‐level community structure and on ecosystem functioning and highlight the value of collaborative, systematic sampling effort., We are grateful for data from the Greenland Ecosystem Monitoring Programme provided by the Department of Bioscience, Aarhus University, Denmark, in collaboration with Greenland Institute of Natural Resources, Nuuk, Greenland, and the Department of Biology, University of Copenhagen, Denmark, and data from the Greenland Ecosystem Monitoring Programme provided by Asiaq—Greenland Survey, Nuuk, Greenland. We acknowledge CSC–IT Center for Science Ltd., Espoo, Finland, for the allocation of computational resources. Rósa Erlendsdóttir kindly provided access to the sampling site in Votilækur, Iceland. We thank Marie‐Andree Giroux and Kristen Peck for their assistance in Igloolik and Marjo Kilpinen, Eija Takala and Emma Hakanen for their hard work in the laboratory. We gratefully acknowledge funding from the Academy of Finland (grants 276909, 285803 to T.R., and 276671 to M.V.K.), Nessling Foundation (grants 201700420, 201600034, 201500090 to T.K.), Jane and Aatos Erkko Foundation, Societas pro Fauna et Flora Fennica, The Danish Environmental Protection Agency (N.M.S.), French Polar Institute‐IPEV (program “Interactions 1036” to O.G. and B.S.), INTERACT, Russian Foundation for Basic Research (grant 18‐05‐60261 to A.S. and N.S.), Research Council of Norway (grant 249902/F20), the Natural Sciences and Engineering Research Council of Canada (PGS‐D award to S.K.M., CGS‐D award to A.M.S., Discovery Grant to J.B.), Churchill Northern Studies Centre (Northern Research Fund Award to A.M.S. and M.P.), Canadian Polar Commission (Northern Scientific Training Program Award to D.L.B., K.H.I.D., S.K.M., A.M.S., and M.P.), ArcticNet, Polar Continental Shelf Project, Parks Canada, Fonds Québécois de Recherche ‐ Nature et Technologies, Polar Knowledge Canada, Entomological Society of Canada, University of Guelph. The Icelandic Research Fund (Rannsóknasjóður, grant nr 152468‐051 to I.C.B.). Thanks to PhyloPic contributors Gareth Monger and Melissa Broussard.

Lichens and bryophytes are abundant primary producers in high latitude and high elevation ecosystems, and they play an important role in ecosystem processes such as decomposition and nutrient cycling. Despite their importance, little is known about the decomposability of lichens and bryophytes either among or within species, at the whole community level, or how this decomposability is affected by their functional traits. Here, we studied decomposability of lichens and bryophytes at the community-level and individual species-level (using 21 species and genera) collected from an elevational gradient in alpine Norway. In order to isolate the elevation effect on litter quality, we used a standardized laboratory bioassay to measure decomposability. In contrast to our expectations, we found that community-level decomposability of lichens and bryophytes increased with elevation and thus decreasing temperature. In contrast, phosphorus release from the litter decreased with elevation while nitrogen release was unresponsive. Decomposability was explained by nutrient concentrations, litter pH and primary producer group identity (lichens versus bryophytes) at both the individual species and community levels. Species turnover (changes in species composition and abundance) was the main driver of decomposability across elevation at the community level, despite some of the traits explaining decomposability showing high intraspecific variability. Our study highlights the importance of among-species variation in determining lichen and bryophyte decomposability. Further, the higher decomposability that we found for higher elevations suggests that global warming might result in a shift towards slower decomposable lichen and bryophyte species.

La acción COST NNEXT ha financiado el coste para publicar este artículo en abierto, por lo que aparece en la web de la revista como acceso abierto: https://onlinelibrary.wiley.com/doi/full/10.1111/brv.12511, Non-native tree (NNT) species have been transported worldwide to create or enhance services that are fundamental for human well-being, such as timber provision, erosion control or ornamental value; yet NNTs can also produce undesired effects, such as fire proneness or pollen allergenicity. Despite the variety of effects that NNTs have on multiple ecosystem services, a global quantitative assessment of their costs and benefits is still lacking. Such information is critical for decision-making, management and sustainable exploitation of NNTs. We present here a global assessment of NNT effects on the three main categories of ecosystem services, including regulating (RES), provisioning (PES) and cultural services (CES), and on an ecosystem disservice (EDS), i.e. pollen allergenicity. By searching the scientific literature, country forestry reports, and social media, we compiled a global data set of 1683 case studies from over 125 NNT species, covering 44 countries, all continents but Antarctica, and seven biomes. Using different meta-analysis techniques, we found that, while NNTs increase most RES (e.g. climate regulation, soil erosion control, fertility and formation), they decrease PES (e.g. NNTs contribute less than native trees to global timber provision). Also, they have different effects on CES (e.g. increase aesthetic values but decrease scientific interest), and no effect on the EDS considered. NNT effects on each ecosystem (dis)service showed a strong context dependency, varying across NNT types, biomes and socio-economic conditions. For instance, some RES are increased more by NNTs able to fix atmospheric nitrogen, and when the ecosystem is located in low-latitude biomes; some CES are increased more by NNTs in less-wealthy countries or in countries with higher gross domestic products. The effects of NNTs on several ecosystem (dis)services exhibited some synergies (e.g. among soil fertility, soil formation and climate regulation or between aesthetic values and pollen allergenicity), but also trade-offs (e.g. between fire regulation and soil erosion control). Our analyses provide a quantitative understanding of the complex synergies, trade-offs and context dependencies involved for the effects of NNTs that is essential for attaining a sustained provision of ecosystem services., Ministerio de Ciencia y Competitividad, Comunidad de Madrid

© 2020 The Authors. Global change experiments such as experimental warming and nutrient addition strongly affect the structure and functioning of high latitude and altitude ecosystems. However, it is often unknown to what extend such effects are permanent or whether changes persist after environmental conditions return to pre-treatment levels. In this study, we assess the legacy effects of temperature manipulation and nutrient addition experiments on alpine soil micro-arthropod (i.e., Collembola and Oribatida) communities nine years after the treatments were discontinued. Treatment effects on the vegetation were still detectable six years after cessation, although grazing increased the recovery rate. Because micro-arthropods are often closely associated with vegetation, we expected to find that treatment effects on Collembola and Oribatida abundance and species composition persisted to date, reflecting plant community dynamics. Also, we expected large-bodied, drought-resistant Collembola species that live on top of the soil to show less strong legacy effects. We did not find legacy effects of environmental treatments on Collembola and Mesostigmata in terms of abundance. However, we found persistent changes in community composition of Collembola and Oribatida, suggesting treatment effects persist to date. The generalist Folsomia quadrioculata was the most responsive Collembola species to initial treatments, most likely due to its variable life-history strategy. Although its abundance recovered, F. quadrioculata remained dominant in Collembola communities after cessation of the treatments. Grazing affected community composition of both Collembola and Oribatida, but we did not find grazing to reduce legacy effects on micro-arthropod as it did for vegetation. We therefore conclude that the environmental treatments had only temporary effects on micro-arthropods in terms of overall abundance, but that effects on individual species and therefore species composition may be long-lasting and less predictable. The study was designed by Johan Asplund, Juha M. Alatalo, and Kari Klanderud. Field work was performed by Ruben Erik Roos, Johan Asplund, Kari Klanderud, and Tone Birkemoe. Peter Ľuptáčik and Natália Raschmanová identified soil micro‐arthropods for 2016. Statistical analyses were performed by Ruben Erik Roos and Siri Lie Olsen. All co‐authors contributed to manuscript revisions and agree with the final version. This study was funded by Carl Tryggers stiftelse för vetenskaplig forskning through a grant to Juha M. Alatalo and a grant from the Research Council of Norway (249902) to Johan Asplund. We thank Sigmund Hågvar for sharing his original data, comments and feedback, Hans Cornelissen and Stef Bokhorst for useful discussions, and Matty Berg for sharing data from his personal Collembola database. Mari Steinert, Ross Wetherbee, Mahdieh Tourani, and Richard Bischof were of great help for discussions on the statistical analyses. We thank the Finse Alpine Research Center and Erika Leslie for hospitality during fieldwork and Kristel van Zuijlen for assistance in the field.

Across environmental gradients, community‐level functional traits of plants can change due to species turnover, intraspecific variation and their covariation. Studies on vascular plants suggest that species turnover is the main driver of trait variation across gradients, although intraspecific variation can also be important. However, there is limited knowledge about whether this holds for non‐vascular primary producers such as lichens and bryophytes. We hypothesized that intraspecific variation is more important for non‐vascular than for vascular primary producers because they lack specialized structures to maintain homeostasis and should therefore be more responsive to extrinsic factors. To assess the relative importance of species turnover versus intraspecific variation for vascular plants, lichens and bryophytes, we estimated species abundance and measured chemical (tissue nitrogen (N) and phosphorous (P) content, N:P ratio and pH) and non‐chemical (specific leaf or thallus area, dry matter content and water holding capacity) functional traits along an elevational gradient in alpine southern Norway. We calculated community‐weighted mean traits and quantified the relative contribution of species turnover, intraspecific variation and their covariation to total trait variation across the gradient. We found mixed support for our hypothesis: the contribution of intraspecific variation to total trait variation for N and N:P was higher in lichens than in vascular plants and bryophytes, but in general the contribution of intraspecific variation differed among functional traits and producer groups. Nutrient variables (N, P and N:P) were significantly impacted by intraspecific variation for vascular plants and lichens but not for bryophytes. Non‐chemical traits and pH were mainly driven by species turnover effects in all primary producer groups. Our results highlight that while nearly all studies on primary producer trait variation across environments have focused on vascular plants, trait variation of other largely neglected but ecologically important producer groups, such as lichens and bryophytes, may show very different responses to the same environmental factors. In order to fully understand how future environmental changes impact on community‐ and ecosystem‐level processes, traits of primary producers other than vascular plants—and their within‐species variation—need to be considered in systems where these groups are abundant. A free Plain Language Summary can be found within the Supporting Information of this article. Published version Norges Forskningsråd, Grant/Award Number: 249902/F20

The Arctic is warming twice as fast as the rest of the planet, leading to rapid changes in species composition and plant functional trait variation. Landscape-level maps of vegetation composition and trait distributions are required to expand spatially-limited plot studies, overcome sampling biases associated with the most accessible research areas, and create baselines from which to monitor environmental change. Unmanned aerial vehicles (UAVs) have emerged as a low-cost method to generate high-resolution imagery and bridge the gap between fine-scale field studies and lower resolution satellite analyses. Here we used field spectroscopy data (400–2500 nm) and UAV multispectral imagery to test spectral methods of species identification and plant water and chemistry retrieval near Longyearbyen, Svalbard. Using the field spectroscopy data and Random Forest analysis, we were able to distinguish eight common High Arctic plant tundra species with 74% accuracy. Using partial least squares regression (PLSR), we were able to predict corresponding water, nitrogen, phosphorus and C:N values (r 2 = 0.61–0.88, RMSEmean = 12%–64%). We developed analogous models using UAV imagery (five bands: Blue, Green, Red, Red Edge and Near-Infrared) and scaled up the results across a 450 m long nutrient gradient located underneath a seabird colony. At the UAV level, we were able to map three plant functional groups (mosses, graminoids and dwarf shrubs) at 72% accuracy and generate maps of plant chemistry. Our maps show a clear marine-derived fertility gradient, mediated by geomorphology. We used the UAV results to explore two methods of upscaling plant water content to the wider landscape using Sentinel-2A imagery. Our results are pertinent for high resolution, low-cost mapping of the Arctic.

We studied the effects of different mat-forming lichens on microclimate and litter decomposition in an alpine ecosystem where lichens constitute a major part of the vegetation. We used four lichens with contrasting colour and water-holding capacity. We recorded soil temperature and moisture, and decomposition rate of plant litter under the different lichen mats and in bare soil. While soil temperature and freeze-thaw cycles were reduced under all lichen mats compared to bare soil, Cladonia rangiferina/Cladonia stygia insulated stronger than other lichens. Litter decomposition was faster under Flavocetraria nivalis than under Alectoria ochroleuca, but this was not related to lichen traits or microclimate. We conclude that insulation by lichen mats is stronger with higher water-holding capacity, and we suggest that microbial communities associated with different lichens have a larger effect on litter decomposition than soil conditions. Our findings highlight the importance of mat-forming lichens on microclimate and ecosystem processes in alpine environments.