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Caroline Greiser

About me

mobil: +46 764152485

I am a landscape ecologist with a PhD in Plant Ecology (2020, Stockholm Uni). My research circles around climate at organism-relevant scales, or so-called microclimate. I measure, model and map temperatures near the ground and investigate how this small-scale variation affects ground-dwelling plants and animals. I have worked with mosses, lichens, vascular plants and insects – mainly in the Swedish forest or agricultural landscape.  

Keywords: microclimate, microrefugia, stepping stones, boreal forests, insects, bryophytes, understory biodiversity, climate change


Currently, I am funded by a FORMAS mobility grant for early career researchers (2022-2025) to look at forest microclimate buffering – the air conditioning effect of the forest – and how it is driven and limited by soil water. Within this project, I work with colleagues from SLU in Umeå and Uppsala, as well as researchers from Czech Republic, Finland, Norway, Belgium and France. The goal is to pool our datasets and find generalities and differences in European forests when it comes to forest microclimate buffering.

In the context of this research, I also collaborate with Rob Lewis from Norway in the project "Understanding the role and interplay of forest microclimates for successfully balancing productivity and biodiversity among Nordic forest landscapes (ForestMicroClim)" - funded by SNS.

Additionally, I collaborate with Philipp Lehmann from Greifswald University (Germany), modelling microclimate on very fine scales in order to feed ecophysiological models of the green-veined white butterfly (Pieris napi). In this exciting project, we integrate the non-linear thermal responses of developmental rate over temperature data on very high spatial and temporal resolution (meters and hours) and explore the effects of a natural microclimate landscape on summer phenology, emergence synchrony and voltinism (nr. of generations) of a butterfly population. 

Together we also explore the role of spruce bark beetles (Ips typographus) as a microclimate engineer. This project is funded by the Lamm foundation

I am a steering committee member of SoilTemp, a global network and database for near-ground microclimate and biodiversity data. 

Here you find a PDF of my PhD thesis.

Research projects


A selection from Stockholm University publication database

  • Monthly microclimate models in a managed boreal forest landscape

    2018. Caroline Greiser (et al.). Agricultural and Forest Meteorology 250-251, 147-158


    The majority of microclimate studies have been done in topographically complex landscapes to quantify and predict how near-ground temperatures vary as a function of terrain properties. However, in forests understory temperatures can be strongly influenced also by vegetation. We quantified the relative influence of vegetation features and physiography (topography and moisture-related variables) on understory temperatures in managed boreal forests in central Sweden. We used a multivariate regression approach to relate near-ground temperature of 203 loggers over the snow-free seasons in an area of ∼16,000 km2 to remotely sensed and on-site measured variables of forest structure and physiography. We produced climate grids of monthly minimum and maximum temperatures at 25 m resolution by using only remotely sensed and mapped predictors. The quality and predictions of the models containing only remotely sensed predictors (MAP models) were compared with the models containing also on-site measured predictors (OS models). Our data suggest that during the warm season, where landscape microclimate variability is largest, canopy cover and basal area were the most important microclimatic drivers for both minimum and maximum temperatures, while physiographic drivers (mainly elevation) dominated maximum temperatures during autumn and early winter. The MAP models were able to reproduce findings from the OS models but tended to underestimate high and overestimate low temperatures. Including important microclimatic drivers, particularly soil moisture, that are yet lacking in a mapped form should improve the microclimate maps. Because of the dynamic nature of managed forests, continuous updates of mapped forest structure parameters are needed to accurately predict temperatures. Our results suggest that forest management (e.g. stand size, structure and composition) and conservation may play a key role in amplifying or impeding the effects of climate-forcing factors on near-ground temperature and may locally modify the impact of global warming.

    Read more about Monthly microclimate models in a managed boreal forest landscape
  • Hiding from the climate

    2020. Caroline Greiser (et al.). Global Change Biology 26 (2), 471-483


    Climate warming is likely to shift the range margins of species poleward, but fine-scale temperature differences near the ground (microclimates) may modify these range shifts. For example, cold-adapted species may survive in microrefugia when the climate gets warmer. However, it is still largely unknown to what extent cold microclimates govern the local persistence of populations at their warm range margin. We located 99 microrefugia, defined as sites with edge populations of 12 widespread boreal forest understory species (vascular plants, mosses, liverworts and lichens) in an area of ca. 24,000 km(2) along the species' southern range margin in central Sweden. Within each population, a logger measured temperature eight times per day during one full year. Using univariate and multivariate analyses, we examined the differences of the populations' microclimates with the mean and range of microclimates in the landscape, and identified the typical climate, vegetation and topographic features of these habitats. Comparison sites were drawn from another logger data set (n = 110), and from high-resolution microclimate maps. The microrefugia were mainly places characterized by lower summer and autumn maximum temperatures, late snow melt dates and high climate stability. Microrefugia also had higher forest basal area and lower solar radiation in spring and autumn than the landscape average. Although there were common trends across northern species in how microrefugia differed from the landscape average, there were also interspecific differences and some species contributed more than others to the overall results. Our findings provide biologically meaningful criteria to locate and spatially predict potential climate microrefugia in the boreal forest. This opens up the opportunity to protect valuable sites, and adapt forest management, for example, by keeping old-growth forests at topographically shaded sites. These measures may help to mitigate the loss of genetic and species diversity caused by rear-edge contractions in a warmer climate.

    Read more about Hiding from the climate
  • Climate limitation at the cold edge

    2020. Caroline Greiser (et al.). Ecography 43 (5), 637-647


    The role of climate in determining range margins is often studied using species distribution models (SDMs), which are easily applied but have well-known limitations, e.g. due to their correlative nature and colonization and extinction time lags. Transplant experiments can give more direct information on environmental effects, but often cover small spatial and temporal scales. We simultaneously applied a SDM using high-resolution spatial predictors and an integral projection (demographic) model based on a transplant experiment at 58 sites to examine the effects of microclimate, light and soil conditions on the distribution and performance of a forest herb, Lathyrus vernus, at its cold range margin in central Sweden. In the SDM, occurrences were strongly associated with warmer climates. In contrast, only weak effects of climate were detected in the transplant experiment, whereas effects of soil conditions and light dominated. The higher contribution of climate in the SDM is likely a result from its correlation with soil quality, forest type and potentially historic land use, which were unaccounted for in the model. Predicted habitat suitability and population growth rate, yielded by the two approaches, were not correlated across the transplant sites. We argue that the ranking of site habitat suitability is probably more reliable in the transplant experiment than in the SDM because predictors in the former better describe understory conditions, but that ranking might vary among years, e.g. due to differences in climate. Our results suggest that L. vernus is limited by soil and light rather than directly by climate at its northern range edge, where conifers dominate forests and create suboptimal conditions of soil and canopy-penetrating light. A general implication of our study is that to better understand how climate change influences range dynamics, we should not only strive to improve existing approaches but also to use multiple approaches in concert.

    Read more about Climate limitation at the cold edge
  • Warm range margin of boreal bryophytes and lichens not directly limited by temperatures

    2021. Caroline Greiser (et al.). Journal of Ecology 109 (10), 3724-3736


    Species at their warm range margin are potentially threatened by higher temperatures, but may persist in microrefugia. Whether such microsites occur due to more suitable microclimate or due to lower biotic pressure from, for example competitive species, is still not fully resolved. We examined whether boreal bryophytes and lichens show signs of direct climate limitation, that is whether they perform better in cold and/or humid microclimates at their warm range margin. We transplanted a moss, a liverwort and a lichen to 58 boreal forest sites with different microclimates at the species' southern range margin in central Sweden. Species were grown in garden soil patches to control the effects of competitive exclusion and soil quality. We followed the transplanted species over three growing seasons (2016-2018) and modelled growth and vitality for each species as a function of subcanopy temperature, soil moisture, air humidity and forest type. In 2018, we also recorded the cover of other plants having recolonized the garden soil patches and modelled this potential future competition with the same environmental variables plus litter. Species performance increased with warmer temperatures, which was often conditional on high soil moisture, and at sites with more conifers. Soil moisture had a positive effect, especially on the moss in the last year 2018, when the growing season was exceptionally hot and dry. The lichen was mostly affected by gastropod grazing. Recolonization of other plants was also faster at warmer and moister sites. The results indicate that competition, herbivory, shading leaf litter and water scarcity might be more important than the direct effects of temperature for performance at the species' warm range margin. Synthesis. In a transplant experiment with three boreal understorey species, we did not find signs of direct temperature limitation towards the south. Forest microrefugia, that is habitats where these species could persist regional warming, may instead be sites with fewer competitors and enemies, and with sufficient moisture and more conifers in the overstorey.

    Read more about Warm range margin of boreal bryophytes and lichens not directly limited by temperatures
  • Forest microclimates and climate change

    2021. Pieter De Frenne (et al.). Global Change Biology 27 (11), 2279-2297


    Forest microclimates contrast strongly with the climate outside forests. To fully understand and better predict how forests' biodiversity and functions relate to climate and climate change, microclimates need to be integrated into ecological research. Despite the potentially broad impact of microclimates on the response of forest ecosystems to global change, our understanding of how microclimates within and below tree canopies modulate biotic responses to global change at the species, community and ecosystem level is still limited. Here, we review how spatial and temporal variation in forest microclimates result from an interplay of forest features, local water balance, topography and landscape composition. We first stress and exemplify the importance of considering forest microclimates to understand variation in biodiversity and ecosystem functions across forest landscapes. Next, we explain how macroclimate warming (of the free atmosphere) can affect microclimates, and vice versa, via interactions with land-use changes across different biomes. Finally, we perform a priority ranking of future research avenues at the interface of microclimate ecology and global change biology, with a specific focus on three key themes: (1) disentangling the abiotic and biotic drivers and feedbacks of forest microclimates; (2) global and regional mapping and predictions of forest microclimates; and (3) the impacts of microclimate on forest biodiversity and ecosystem functioning in the face of climate change. The availability of microclimatic data will significantly increase in the coming decades, characterizing climate variability at unprecedented spatial and temporal scales relevant to biological processes in forests. This will revolutionize our understanding of the dynamics, drivers and implications of forest microclimates on biodiversity and ecological functions, and the impacts of global changes. In order to support the sustainable use of forests and to secure their biodiversity and ecosystem services for future generations, microclimates cannot be ignored.

    Read more about Forest microclimates and climate change
  • Climate adaptation of biodiversity conservation in managed forest landscapes

    2022. Kristoffer Hylander (et al.). Conservation Biology 36 (3)


    Conservation of biodiversity in managed forest landscapes needs to be complemented with new approaches given the threat from rapid climate change. Most frameworks for adaptation of biodiversity conservation to climate change include two major strategies. The first is the resistance strategy, which focuses on actions to increase the capacity of species and communities to resist change. The second is the transformation strategy and includes actions that ease the transformation of communities to a set of species that are well adapted to the novel environmental conditions. We suggest a number of concrete actions policy makers and managers can take. Under the resistance strategy, five tools are introduced, including: identifying and protecting forest climate refugia with cold-favored species; reducing the effects of drought by protecting the hydrological network; and actively removing competitors when they threaten cold-favored species. Under the transformation strategy, we suggest three tools, including: enhancing conditions for forest species favored by the new climate, but currently disfavored by forest management, by planting them at suitable sites outside their main range; and increasing connectivity across the landscape to enhance the expansion of warm-favored species to sites that have become suitable. Finally, we suggest applying a landscape perspective and simultaneously managing for both retreating and expanding species. The two different strategies (resistance and transformation) should be seen as complementary ways to maintain a rich biodiversity in future forest ecosystems. 

    Read more about Climate adaptation of biodiversity conservation in managed forest landscapes
  • Microclimatic variation affects developmental phenology, synchrony and voltinism in an insect population

    2022. Caroline Greiser (et al.). Functional Ecology 36 (12), 3036-3048

    1. Temperature influences the rate of most biological processes. Nonlinearities in the thermal reaction norms of such processes complicate intuitive predictions of how ectothermic organisms respond to naturally fluctuating temperatures, and by extension, to climate warming. Additionally, organisms developing close to the ground experience a highly variable microclimate landscape that often is poorly captured by coarse standard climate data.
    2. Using a butterfly population in central Sweden as a model, we quantified the consequences of small-scale temperature variation on phenology, emergence synchrony and number of annual reproductive cycles (voltinism). By combining empirical microclimate and thermal performance data, we project development of individual green-veined white butterflies (Pieris napi) across 110 sites in an exceptionally high-resolved natural microclimate landscape.
    3. We demonstrate that differences among microclimates just meters apart can have large impacts on the rate of development and emergence synchrony of neighbouring butterflies. However, when considering the full development from egg to adult, these temporal differences were reduced in some scenarios, due to negative correlations in development times among life stages. The negative correlations were caused by temperatures at some sites beginning to exceed the optimum for development as the season progressed. Indeed, which sites were optimal for fast development could change across the lifetimes of individual butterflies, that is, ‘fast’ sites could become ‘slow’ sites. Thus, from a thermal point of view, there seem to be no consistently optimal microsites. Importantly, the fast sites were not always the warmest sites. We showed that such unintuitive effects could play an important role in the regulation of phenological synchrony and voltinism in insects, as most sites consistently favoured two generations. The results were generally robust across years and three different egg-laying dates.
    4. Using high-resolved empirical climate data on organism-relevant temporal and spatial scales and considering nonlinear responses to temperature, we demonstrated the large and unintuitive population-level consequences of locally and temporarily high temperatures. We suggest to—whenever possible—incorporate species- and life stage-specific nonlinear responses to temperature when studying the effects of natural microclimate variation and climate change on organisms.
    Read more about Microclimatic variation affects developmental phenology, synchrony and voltinism in an insect population
  • Maintaining forest cover to enhance temperature buffering under future climate change

    2022. Emiel De Lombaerde (et al.). Science of the Total Environment 810


    Forest canopies buffer macroclimatic temperature fluctuations. However, we do not know if and how the capacity of canopies to buffer understorey temperature will change with accelerating climate change. Here we map the difference (offset) between temperatures inside and outside forests in the recent past and project these into the future in boreal, temperate and tropical forests. Using linear mixed-effect models, we combined a global database of 714 paired time series of temperatures (mean, minimum and maximum) measured inside forests vs. in nearby open habitats with maps of macroclimate, topography and forest cover to hindcast past (1970–2000) and to project future (2060–2080) temperature differences between free-air temperatures and sub-canopy microclimates. For all tested future climate scenarios, we project that the difference between maximum temperatures inside and outside forests across the globe will increase (i.e. result in stronger cooling in forests), on average during 2060–2080, by 0.27 ± 0.16 °C (RCP2.6) and 0.60 ± 0.14 °C (RCP8.5) due to macroclimate changes. This suggests that extremely hot temperatures under forest canopies will, on average, warm less than outside forests as macroclimate warms. This knowledge is of utmost importance as it suggests that forest microclimates will warm at a slower rate than non-forested areas, assuming that forest cover is maintained. Species adapted to colder growing conditions may thus find shelter and survive longer than anticipated at a given forest site. This highlights the potential role of forests as a whole as microrefugia for biodiversity under future climate change.

    Read more about Maintaining forest cover to enhance temperature buffering under future climate change

Show all publications by Caroline Greiser at Stockholm University