The role of microclimate for the

performance and distribution of

forest plants

Carl Johan Dahlberg

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© C. Johan Dahlberg, Stockholm University 2016 Front cover illustration: Niklas Lönnell ISBN 978-91-7649-423-3 Printed in Sweden by Holmbergs, Malmö 2016 Distributor: Department of Ecology, Environment and Plant Sciences, Stockholm University

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To Sara and Linnéa

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Abstract Microclimatic gradients may have large influence on individual vital rates and population growth rates of species, and limit their distributions. Therefore, I focused on the influence of microclimate on individual performance and distribution of species. Further, I examined differences in how microclimate affect species with contrasting distributions or different ecophysiological traits, and populations within species. More specifically, I investigated the performance of northern and southern distributed forest bryophytes that were transplanted across microclimatic gradients, and the timing of vegetative and reproductive development among northern, marginal and more southern populations of a forest herb in a common garden. Also, I compared the landscape and continental distributions across forest bryophytes and vascular plants and, thus, their distribution limiting factors at different spatial scales. Lastly, I examined the population dynamics across microclimatic gradients of transplants from northern and southern populations of a forest moss. The effects of microclimatic conditions on performance differed among bryophytes with contrasting distributions. There were no clear differences between northern and southern populations in the timing of development of a forest herb or in the population dynamics of a moss. However, within each region there was a differentiation of the forest herb populations, related to variation in local climatic conditions and in the south also to proportion of deciduous trees. The continental distributions of species were reflected in their landscape distributions and vice versa, in terms of their occurrence optima for climatic variables. The variation in landscape climatic optima was, however, larger than predicted, which limit the precision for predictions of microrefugia. Probably, the distributions of vascular plants were more affected by temperature than the distributions of bryophytes. Bryophytes are sensitive to moisture conditions, which was demonstrated by a correlation between evaporation and the population growth rate of a forest moss. We might be able to predict species’ landscape scale distributions by linking microclimatic conditions to their population growth rates, via their vital rates, and infer larger scale distribution patterns. Keywords: microclimate, spatial scales, continental, landscape, distribution patterns, distribution limits, phenology, vegetative development, reproductive development, performance, vital rates, population growth rate, vascular plants, bryophytes, distribution modelling

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List of Papers This thesis is based on the following papers, which are referred to by their roman numerals: I. Dahlberg, C. J., Ehrlén, J. Hylander, K. 2014. Performance of forest bryophytes with different geographical distributions transplanted across a topographically heterogeneous landscape. PLoS ONE 11/2014. DOI: 10.1371/journal.pone.0112943 II. Dahlberg, C. J.*, Fogelström, E.*, Hylander, K., Meineri, E., Ehrlén, J. Population differentiation in timing of development in a forest herb associated with local climate and canopy closure. Manuscript III. Dahlberg, C. J., Ehrlén, J. Meineri, E. Hylander, K. Plant landscape climatic optima correlate with their continental range optima. Manuscript IV. Dahlberg, C. J., Ehrlén, J. Hylander, K. Population dynamics of moss transplants across microclimatic gradients. Manuscript The contributors to the papers were: I. Conceived and designed the experiments: CJD JE KH. Performed the experiments: CJD. Analyzed the data: CJD. Led the writing: CJD II. Conceived and designed the experiments: CJD EF KH JE. Performed the experiments: CJD EF. Analyzed the data: CJD EF. Led the writing: CJD EF. *these authors contributed equally to paper (II) III. Conceived and designed the experiments: CJD JE KH. Data collecting: CJD. Analyzed the data: CJD EM. Led the writing: CJD. IV. Conceived and designed the experiments: CJD JE KH. Performed the experiments: CJD. Analyzed the data: CJD. Led the writing: CJD.

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Contents

Introduction……………………………………………………………. 9 Distributions and microclimate………..……………………....... 9 Responses to microclimate……………………………………... 11 Aims of the thesis……………………………………………………… 13 Material and methods………………………………………………….. 14 Study areas……………………………………………………… 14 Northern region………………………………………….. 14 Southern region…………………………………………... 16 Continental region……………………………………….. 17 Study systems…………………………………………………… 17 Bryophytes……………………………………………….. 17 Vascular plants……………………………………………19 Performance of forest mosses across microclimatic gradients (Paper I)………...……………………………….……. 21 Responses of a forest herb to local climate and canopy cover (Paper II)…...………..……………………………….…... 23 Landscape and continental distributions of plants related to climate (Paper III)…………………...……………………….. 26

Population dynamics of a forest moss across microclimatic gradients (Paper IV)…………………………………………….. 28

Results and Discussion………………………………………………… 30 Concluding remarks…………………………………………………… 39 Acknowledgements……………………………………………... 40 References……………………………………………………………... 40 Svensk sammanfattning……………………………………………….. 51 Tack/Acknowledgements……………………………………………… 57

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Introduction

Distributions and microclimate Species are limited in their distributions by environmental conditions (abiotic and biotic factors), their dispersal capacities and physical barriers. Environmental conditions influence their population vital rates, i.e. their performance in terms of growth, survival and reproduction of individuals. Vital rates set the number of births and deaths and, along with immigrants and emigrants, determine population growth rates and species distributions (Gaston, 2003; Gaston, 2009; Ehrlén & Morris, 2015). However, species distribution models often assume that occurrences reflect populations with positive growth rates (e.g. Guisan & Thuiller, 2005; Austin & Van Niel, 2011), i.e. places with environmental conditions within the species’ niches (e.g. Maguire, 1973; Hutchinson, 1978; Holt, 2009), without considering population dynamics. This project is focused on the climatic impact on species distributions, mainly at a local scale through its effects on population vital rates and growth, in order to increase our knowledge on how climatic factors influence species distributions. Climate, and temperature in particular, has often been regarded as the single most important factor for species distributions (Hutchins, 1947; Gaston, 2003). There is ample evidence for how climate regulate species distributions. For example, from plant fossils and seeds we have learnt about species range shifts during historical cold glacials and warmer interglacials (e.g. Birks & Willis, 2008). We often correlate species distributions to variation in temperature or precipitation at rougher scales, and from these correlations we also predict species future distributions under climatic change scenarios (Parmesan, 2006). To learn more about how climate regulate species distributions, I examined the microclimate that is experienced by individuals in situ. Microclimate can be defined as the sum of climatic influences from movements of the free atmosphere and local climatic-forcing factors, such as terrain and vegetation (e.g. Dobrowski, 2011; Hampe & Jump, 2011). I

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investigated how microclimate affect individual performance and species distributions at, for example, their distribution limits. At a trailing range margin, there are certain sites or refugia with favourable environmental conditions where populations can survive, until the main range advances once again (Fig 1; Ashcroft, 2010). In topographic heterogeneous landscapes with large microclimatic variation, such patches with favourable environmental conditions are often rather small. Their climatic trends can be decoupled from the average regional climatic trend due to the local topography and vegetation (Dobrowski, 2011; Hampe & Jump, 2011). For example, may slope direction influence incoming solar radiation which in turn influence the diurnal maximum temperature (Dahlberg et al., 2014). In recent years, the importance of microclimate for species distributions have been emphasized (e.g. Scherrer & Körner, 2011; Ashcroft & Gollan, 2012). Topographic heterogeneous landscapes might buffer the impact of climate change on species range shifts (Loarie et al., 2009). We need to increase our knowledge about how microclimate regulate species distributions, in order to refine predictions of current and future distribution patterns.

Figure 1. Hypothetical northern and southern range margins with fragmented populations beyond the main distribution range. These may correspond to refugia or stepping stones if the margin is a trailing or leading edge, respectively.

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Responses to microclimate During climatic change, populations may respond through adaptive genetic evolution or phenotypic plasticity to the new climatic conditions and, thereby, survive. Populations may also track the trailing climate through migration or dispersal, or they may go extinct (Franks et al., 2014). I examined population responses that occur genetically or through plasticity. Genetic adaptation occur through natural selection that increase the performance and fitness of individuals and, thus, population growth rates (cf. Linhart & Grant, 1996; Hendry & Kinnison, 2001; Eckhart et al., 2011). This differs from plastic response of individuals to fluctuating climatic conditions within a generation, which do not include genetic change (Bradshaw, 1965). A large plastic capacity of individuals may increase the resilience of populations to climatic change. However, if climatic change continues over generations and exceed the niches of species, phenotypic plasticity is not enough to secure the population survival. Still, the species may survive through selective adaptation (Hampe & Jump, 2011). Relatively few studies concern how adaptation to climatic conditions influence population vital rates and population growth (e.g. Doak & Morris, 2010; Buckley & Kingsolver, 2012; Stevens & Latimer, 2015), although local adaptation within species is generally regarded to be common in nature (Hendry & Kinnison, 2001). Such studies must either span several generation times or population studies across spatial scales. Studies of variation in population vital rates and growth, and differences among populations, across temporal fluctuations in climatic conditions are slightly more common (e.g. Altwegg et al., 2005; Griffith & Loik, 2010; Sletvold et al., 2013). I chose to compare populations of plants from different geographical regions, both in common garden and transplant experiments. In common garden experiments, we can identify population differentiation among populations when grown under the same climatic conditions. In transplant experiments, we can identify differences among populations, since they are transplanted at the same sites. Also, we can study plastic responses of the individuals, population vital rates and population growth rates across the microclimatic gradients of the

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transplant sites. If the genetic differentiation in phenotypic traits act in the same direction as the plastic response to the environmental conditions, the phenotypic variation follow a co-gradient pattern. Similarly, if the plastic response oppose the genetic differentiation, the phenotypic variation follow a counter-gradient pattern (Conover & Schultz, 1995; Conover et al., 2009). To further complicate the picture, there may be differences in plastic responses to environmental conditions between individuals from different regions, i.e. there is an adaptation of plastic response (Lonsdale & Levinton, 1985; Toftegaard et al., 2015).

According to the AASL-hypothesis, stressful conditions at species range margins limit their occurrence by exceeding the physiological tolerance of individuals (Normand et al., 2009). Subsequently, local adaptation have been argued to be most common in populations at range margins, since they are often situated in sub-optimal environmental conditions (e.g. Lesica & Allendorf, 1995; Galloway & Fenster, 2000; Normand et al., 2009). The colder conditions at higher altitudes or latitudes have often been regarded to limit distributions for temperate species, while biotic factors such as inter-specific competitions may limit their distributions margins at lower altitudes and latitudes (Normand et al., 2009; Pellissier et al., 2013). However, this is a simplified viewpoint, since for example drought may be more pronounced at lower altitudes and latitudes, and thereby exert increased stress and limit distributions (cf. Gimenez-Benavides et al., 2008; Ågren & Schemske, 2012). Also, it is probable that the limiting factors, by affecting population vital rates and growth rates, differ between species or species groups with different physiological traits (Davis et al., 1998; Ehrlén & Morris, 2015). Thus, it is important to investigate how microclimate influence the population dynamics and limit distributions across different climatic conditions for many different species and species groups. We still have little knowledge on this matter, although it may help us to better predict the climatic change impact on species distributions.

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Aims of the thesis The main objective of this thesis was to study the influence of microclimate on individual performance and distribution of forest plants, and how performance and distribution are interconnected. Further, I examined differences in how microclimate influence species with different ecophysiological traits (bryophytes and vascular plants), species with contrasting distributions, and populations within species.

Specific objects of the thesis were: • To investigate how the microclimate influence the growth of

one relatively northern (Barbilophozia lycopodioides) and two southern (Eurhynchium angustirete and Herzogiella seligeri) distributed forest bryophytes, by measuring the growth of transplants across a topographically heterogeneous landscape in the main range of the northern species and at the range margins of the southern species.

• To assess the impact of local climate and canopy cover on the population differentiation in development time among relatively southern, main range and northern, marginal distributed populations of a forest herb (Lathyrus vernus).

• To explore how the local climate regulate the landscape distributions of forest vascular plants and bryophytes as compared to how climate influence their continental distributions, and if there are differences between the species groups in how climate influence their distributions.

• To examine the impact of microclimatic conditions on the responses of vital rates, shoot growth and population growth rate, and the genetic differentiation in population dynamics, among relatively northern and more southern populations of a forest bryophyte (Hylocomiastrum umbratum).

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Material and methods Study areas All four papers comprise study landscapes within central Sweden (Fig 2), which mainly consist of boreal forests in the middle boreal subzone. In paper (I) I compared transplants of bryophyte species that I gathered from central and southern Sweden. In paper (III), I compared species’ distributions in a focal landscape of central Sweden with their European, continental distributions. In paper (II) and (IV), I compared populations from central and southern Sweden.

Figure 2. Study areas of the thesis, and the compared northern and southern populations of Lathyrus vernus in paper II and of Hylocomiastrum umbratum in paper IV. Background overview maps: © Lantmäteriet Gävle 2014 (I2014/00691).

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Northern region The focal landscape used for the transplant experiment and species distributions in paper (I) and paper (III) is situated in the county of Ångermanland (between the latitudes 62°50’ and 63°12’ N), and is 73 km from east to west and 42 km from north to south (Figs 2 & 3). This landscape is different from most other areas of Sweden, concerning its hilly terrain that reaches the Bothnian Sea in the east. The altitudinal range goes from 0 m.a.s.l. by the sea to 470 m.a.s.l in the inland (Fig 3), while the mean temperature is about 15.6°C in July and the annual precipitation 671 mm (by the city of Kramfors;

Swedish Meteorological and Hydrological Institute, 2016). The bedrock consist mainly of gneiss with podzolic soils of sandy/loamy/silty materials. The area belongs mainly to the middle boreal subzone (Sjörs, 1999). The microclimate of the area is varied and therefore suitable for climatic studies. It is mainly influenced by the hilly terrain, the Ångerman River and the Bothnian Sea

Figure 3. Focal landscapes and study sites in Ångermanland and Medelpad for inventories and transplant experiments, presented on a Digital Elevation Model (DEM, 50×50 m) with elevation in meters. Background overview maps: © Lantmäteriet Gävle 2014 (I2014/00691).

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(Vercauteren et al., 2013a; Meineri et al., 2015). Moreover, it is suitable for range margin studies, since many southern and also northern plant species have their northern and southern range limits, respectively, in this landscape (Mascher, 1990; Artdatabanken SLU, 2016).

The northern study landscape of paper (II) comprise 10 sites of population origins (between the latitudes 62°39’ and 63°24’ N), is larger but overlapping with the first landscape (I, III) and have similar terrain, boreal forest types and geological characters. Also, the study landscape used in paper (IV) is overlapping with the landscape described above (I, III) (Fig 3). However, it is smaller with a size of 26×37 km and is located only in the inland in southern Ångermanland and northern Medelpad (between the latitudes 62°47’ and 63°08’ N). In this way, we reduced the climatic influence from the Bothnian Sea and the Ångerman River. The forested, northern origin sites for transplants of paper (IV), are situated in the southern boreal subzone (Fig 2). Southern region For paper (I), the transplant material of the southern distributed bryophytes Herzogiella seligeri and Eurhynchium angustirete was collected at a relatively southern site 330 km south of the transplantation experiment (Erken, Lat 59°50’6.30’’, Long 18°30’15.03’’) in the county of Uppland. This site is located in the hemiboreal subzone (Sjörs, 1999). The tree layer was dominated by Norway spruce Picea abies. The more northern distributed bryophyte Barbilophozia lycopodioides was gathered from a north-facing slope in the county of Ångermanland dominated by P. abies (Latberget; Lat 62°56’4.96’’; Long 17°42’19.74’’), with about 20 days shorter growing season than the southern site (Sjörs, 1999).

In paper (II), the southern study landscape comprising 10 sites of population origins is situated in the counties of Uppland, Södermanland and Östergötland (between the latitudes 58°24’ and 60°45’ N; Fig 2). It differ from the northern region in the relatively flat landscape reaching c. 200 ma.s.l. The bedrock is rather similar with mainly gneiss and granite. However, instead of podzolic soil,

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the southern sites have brown earth soils of both sandy materials and clay. The sites consisted of deciduous forests of the hemiboreal subzone (Sjörs, 1999; SLU, 2014), with agricultural land adjacent to some of them. In July the mean temperature is 16.8 °C, while the annual precipitation reach 515 mm (the city of Stockholm, Swedish Meteorological and Hydrological Institute, 2016).

For paper (IV), I gathered transplant material of H. umbratum at three southern sites in the boreonemoral zone with mixed or deciduous tree layers (Fig 2). They have 20 – 30 days longer growing season (Sjörs, 1999) than the northern origin sites, but rather similar annual precipitation (for the period 1961-1990; Swedish Meteorological and Hydrological Institute, 2016).

Continental region In paper (III), the continental scale study area represent a large part of Europe between the latitudes 27°38’ and 81°48’ N (Fig 2). It covers a large altitudinal gradient from below the sea level to the mountain summits. There are several climatic gradients of the area; from cold climates and long winters in the north and at high altitudes to Mediterranean climate with warm summers and mild winters in the south, and east-west gradients with colder and drier climate in the eastern inlands to milder and humid climates by the Atlantic Sea in the west (Peel et al., 2007; Europe, 2014).

Study systems Bryophytes Bryophytes, including mosses (Fig 4), liverworts and hornworts, evolved from green algae and were the first green plants to colonize land for more than 400 million years ago (Slack, 2011). It is yet uncertain if these three groups are monophyletic or paraphyletic (Qiu et al., 2006; Knoop, 2010; Cox et al., 2014). Bryophytes is a widespread and the second most species rich plant group after angiosperms with c. 16 000 species worldwide (Vanderpoorten &

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Goffinet, 2009; Proctor, 2011; Slack, 2011). They are rather species rich in boreal relative tropical regions as compared to vascular plants (Slack, 2011). Bryophytes have an important role for the function of boreal forests through insulating the ground from temperature extremes and absorbing nutrients from the precipitation (Longton, 1979; Bonan & Shugart, 1989; Økland & Eilertsen, 1993; Økland, 1994).

Bryophytes are suitable for studies on microclimatic change, since their growth is highly dependent on their immediate surroundings due to their ecophysiological traits (e.g. Dahlberg et al., 2014). In contrast to vascular plants, they lack water-conducting roots, protective cuticle and regulative stomata, which means that they cannot regulate the uptake and loss of water, i.e. they are poikilohydric. They grow under moist conditions, and dry out during drought until water availability return. Bryophytes have the ability to survive the drought state, albeit frequent drought and rewetting events can induce physiological stress. Most pleurocarps transport water outside their stems, i.e. they are ectohydric, and lack supporting water-conducting xylem. These physiological traits could explain their small size. Moreover, all bryophytes need water for their reproduction, similarly to the spore-producing vascular plants (Goffinet et al., 2009; Proctor, 2009, 2011).

In paper (I), I studied the two pleurocarpus mosses

Herzogiella seligeri [Brid.] Iwats. and Eurhynchium angustirete [Broth.] T.J. Kop. (Fig 4a), and the liverwort Barbilophozia

Figure 4. Two mosses with a relatively southern distribution in Sweden; a) Eurhynchium angustirete that occurs only south of Ångermanland, flanked by a ruler for size measurements, and b) Buxbaumia viridis that has scattered occurrences in Ångermanland, and c) a relatively more northern distributed moss, Hylocomiastrum umbratum, which is common in Ångermanland.

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lycopodioides [Wallr.] Loeske. The two mosses have a southern distribution in Sweden where they are quite common in relatively warm deciduous forests. They have only scattered occurrences in the boreal region along their northern range margin in central Sweden. The liverwort B. lycopodioides have a more northern, boreal distribution in Sweden where it is common especially in locally cool and moist forests. However, it also occur with more scattered occurrences in southern Sweden (Söderström, 1981; Artdatabanken SLU, 2016). All three species are perennials and can form large dominating carpets at favourable sites. However, the relatively large species E. angustirete and B. lycopodioides grow mostly on the forest floor and can stay for a long time in the same patches, while the smaller H. seligeri mostly is confined to continuously changing patches of dead wood (cf. During, 1979; Cronberg & Darell, 2011).

The study species of the paper (IV), Hylocomiastrum umbratum [Hedw.] Fleisch. (Fig 4c), is a relatively large pleurocarpus moss (Smith, 2004). It has an incomplete circumpolar distribution and prefer suboceanic and oceanic regions (Ratcliffe, 1968; Koponen, 1979). Its main range in Scandinavia is located in maritime parts of Svealand and southern Norrland in Sweden and in southwestern Norway. It has scattered occurrences in Finland and southernmost and northern Sweden (Nitare, 2000; GBIF, 2015). It mostly grow in shaded and moist forest habitats such as ravines and on north-facing slopes (Söderström, 1981; Nitare, 2000). It grow in carpets directly on the forest floor or on logs and boulders, similarly to E. angustirete and B. lycopodioides. Just like Hylocomium splendens, it produces one or several new segments per shoot every autumn which mature in the following autumn, and is therefore relatively easy to examine both regarding shoot growth and growth form (Økland, 1995; Hylander et al., 2002; Hylander, 2005). It can also produce new segments by apical growth and branching following segment breaks (IV).

Vascular plants Vascular plants have dominated the flora during the last 300 million years, including the spore-bearing club mosses, horse-tails and ferns

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and the seed-producing gymnosperms and angiosperms (Bell & Hemsley, 2000; Proctor, 2011). It is the second most species rich organism group after beetles, comprising around 300 000 of species worldwide (Kreft & Jetz, 2007). In the northern hemisphere, they form the circumpolar boreal forests which is the focus study system of this thesis. Boreal forests belong to the largest terrestrial biome on earth (Sanderson et al., 2012), although they contain only a tiny portion of the total vascular plant species. The tree layer in boreal forests is species poor with Norway spruce Picea abies [L.] Karst., Scots pine Pinus sylvestris L. and a few deciduous tree species, whereas the ground layer is richer in vascular plants (Kuusipalo, 1985). Dwarf-shrubs, grasses and herbs (Fig 5) makes up the vegetation classes of the vascular plants ground layer (Hägglund & Lundmark, 1982).

A number of ecophysiological traits could be attributed to the prominent role of vascular plants in many ecosystems. First, they have water-transporting roots and leaves with a protective layer of cuticle and stomata which can control water uptake and loss, i.e. they are homoiohydric. Secondly, they have an internal water-conducting and supporting tissue, the xylem, which means that they are endohydric. Because of their homoiohydric and endohydric states, they can keep water in their tissues and continue to grow, with mechanical support, even under temporarily dry weather. It explain why most vascular plants can grow larger than for example bryophytes. Moreover, the growth in trees and shrubs is supported by lignified xylem (Proctor, 2007, 2011).

In paper (II), we study the forest herb Lathyrus vernus [L.] Bernh. (Fig 5a). It is distributed in a large part of central and eastern Europe (Hultén & Fries, 1986). In Sweden, it occur from the southernmost parts up to the boreal region in central Sweden (Anderberg & Anderberg, 2014). The boreal occurrences are thus only small populations at the northern range margin of the species, and probably remnants from earlier warmer periods (Widén & Schiemann, 2003). It is a rather long-lived species with an average life-span of 44.1 years (Ehrlén & Lehtilä, 2002). It has a subterranean rhizome from which erect shoots of 5 – 40 cm grows

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up in early spring. It develop red-purple flowers which open a few weeks after shoot emergence (Ehrlén, 1995; Ehrlén & Münzbergová, 2009). Flowering phenology and shoot emergence date are correlated. The next year’s shoots and flowers are initiated already before fruit maturation (Sola & Ehrlén, 2007).

Performance of forest mosses across microclimatic gradients (Paper I) I investigated the performance, in terms of growth, vitality and capsule maturation, of the forest bryophytes B. lycopodioides, E. angustirete (Fig 3a) and H. seligeri transplanted at 15 south- and 18 north-facing slopes in the county of Ångermanland in central Sweden (Figs 2 & 3). E. angustirete was transplanted north of its northern range margin, H. seligeri was transplanted at or north of its range margin, and the more northern distributed B. lycopodioides was transplanted in its main range (Artdatabanken SLU, 2016). I related their performance to measurements of both air and ground temperature of the sites. The transplant sites consisted of forests with a mature tree layer of at least 50 years of age. Norway spruce dominated the tree layer at most sites. At each site, I selected a 4 x 4 meter square on mesic soil, away from open ground, younger forest stands, rocky outcrops or streams that could influence the microclimate. I gathered transplant material from two origin sites (see above) in April 2011 by carefully removing patches of E. angustirete and B. lycopodioides from the soil, and pieces of wood with H. seligeri from the logs were it was growing. In April to May

Figure 5. Two herbs with a relatively southern distribution in Sweden, with scattered occurrences in Ångermanland; a) Lathyrus vernus and b) Actaea spicata, and c) a herb with a more northern distribution, Lactuca alpina which is rather common in Ångermanland.

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2011, I removed the vegetation down to the soil and attached transplant material of three patches per species in each square. The patches were about 10 cm in diameter for E. angustirete, 5 cm for B. lycopodioides and 3 cm for H. seligeri. I measured growth during the experimental season (April to October 2011) as transplant size change through photographs of B. lycopodioides and E. angustirete. Vitality at the end of the experimental period was estimated in the field for all three species following the scale: (1) <50% of the transplant was freshly green (vigorous); (2) 50-<95% of the transplant was freshly green; (3) 95% of the transplant was freshly green. Capsule maturation was only measured for H. seligeri by calculating the proportion of capsules that matured during the experimental period.

I measured air and ground temperatures with two ibuttons at each site. The air temperature was measured 1 m above the ground by sheltering the logger in a white plastic cup (Fig 6a). The ground temperature was measured underneath the moss cover by placing the logger inside two small zip-bags. The loggers recorded temperature every half hour during the experimental period. From these measurements, I investigated how four microclimatic variables influenced performance: extreme cold air temperature, mild maximum air temperature, extreme warm air temperature and diurnal ground temperature range. The extreme cold air temperature was calculated as the 5th percentile coldest temperature, while the mild and extreme warm temperatures were calculated as the 5th and 95th percentiles of maximum temperature, respectively. While the 5th percentile coldest temperature and the 95th percentile warmest temperature represent extreme cold and extreme heat, respectively, a high mild maximum temperature (5th percentile of maximum temperature) represent a site were most of the days remained relatively warm. I used percentiles in order to reduce the influence of outliers.

Also, I examined how four other environmental variables affected performance. These were: distance to the sea, distance to open ground, solar radiation and productivity. Solar radiation (direct radiation plus diffuse radiation in kWh/m2) for each site was

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calculated between the 29th of May and the 28th of September of 2011, by using solar radiation tool (spatial analyst extension) on a 50 × 50 meter digital elevation model (DEM) in ArcGIS Desktop 10.0. The shortest distance to the sea and distance to open ground from the site midpoints were measured on topographic maps and on aerial photos in ArcGIS Desktop 10.0. Productivity was estimated by classifying the vegetation within each site according the following scale, which indicate increasing productivity: (1) dwarf-shrubs Calluna vulgaris and Empetrum nigrum; (2) dwarf-shrub Vaccinium vitis-idaea; (3) dwarf-shrub V. myrtillus; (4) low herbs and dwarf-shrubs (V. myrtillus); (5) low herbs or (6) tall herbs (Hägglund & Lundmark, 1982). Responses of a forest herb to local climate and canopy cover (Paper II) We investigated the population differentiation in the timing of vegetative and reproductive development in the forest herb L. vernus

Figure 6. a) A shielded air temperature logger (ibutton) in a boreal forest habitat at one of the sites. b) A plastic cylinder that was used for measurement of evaporation at one of the sites for paper (IV). The cylinder was filled with distilled water. We attached a filter paper at the bottom though which the water could evaporate. The picture also illustrate a plastic mug with an ibutton inside for measurements of humidity and near-ground temperature.

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(Fig 5a). We examined start of development, development time and start of flowering among northern, marginal populations and more southern, central populations (Fig 2), when grown in a common environment. We related the development variables to proportions of deciduous trees and maximum temperatures of the population origin sites. Also, we investigated the relationships among the three development variables.

We gathered fruits from the 10 northern and the 10 southern populations in the summers of 2010 and 2011. Subsequently, we sowed seeds in a greenhouse and examined the plants in a common garden in Stockholm. We retrieved start of development, development time and start of flowering from weekly recordings in a common garden study on 375 plants during spring 2014. For start of development, we used the day when a plant had reached 10% of its final height. We estimated start of flowering by using the size of the largest flower bud in the previous measurement. As development time, we counted the number of days between the start of development and start of flowering.

We estimated the yearly maximum temperature for the site of origin of each population. For the southern region, we predicted maximum temperature for each site by using a linear model based on hourly temperature measurements at 1.5 – 2 m height for ten years (31 December 2013 to 1 January 2014) of 35 stations (5114 km2) managed by the Swedish meteorological and hydrological Institute (SMHI, www.smhi.se). To derive the linear model, we regressed maximum temperature for each station against several topographic variables (cf. Ashcroft & Gollan, 2012). We stepwise, selected elevation and distance to open sea as explanatory variables for the model. To model temperature, we used R version 3.1.1 (R Core Team, 2014). We calculated the physiographic variables from a 50 m grained Digital Elevation Model (Lantmäteriet; the Swedish mapping, cadastral and land registration authority, www.lantmateriet.se) in Arcmap (Version 10.1, Esri, www.esri.com).

For the northern region, we modelled maximum temperature from 1-year hourly temperature measurements during 2011 – 2012

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(Fig 7a; Meineri et al., 2015). These loggers measured temperature at 1 m height in similar forest habitats and microclimatic conditions as the L. vernus populations (Fig 6a), in contrast to the SMHI stations which measured temperature in open areas. We predicted 95th percentile of maximum temperature by using Bayesian Network path models to depict relationships between maximum temperature and topographic variables (Meineri et al., 2015). We estimated the proportion of deciduous trees at the different origin sites by obtaining the proportion of pixels that represented deciduous trees of total grown up forest in infra-red images, covering an area of 50 × 50 m at each site (Lantmäteriet; the Swedish mapping, cadastral and land registration authority, www.lantmateriet.se). We obtained the number of pixels in the program GIMP 2.8.16 (The GIMP team, www.gimp.org).

Figure 7. a) Focal landscape used in paper I and III with temperature logger sites and predicted maximum temperatures ( C) for June 2011 to June 2012. This temperature model was used in paper II and III. b) Focal landscape used in paper I and III with temperature logger sites and predicted growing degree days (GDD5) for June 2011 to June 2012, used in paper III. See Meineri et al. (2015) for methods on the temperature models.

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Landscape and continental distributions of plants related to climate (Paper III) In this study, I investigated if the landscape distributions of species are reflected in their continental distributions and vice versa. More specifically, I examined the correlations between the landscape and continental occurrence optima of 146 forest bryophytes and vascular plants for the climatic variables growing degree days, maximum temperature and minimum temperature. These variables have been suggested to be important for species distributions (e.g. Trivedi et al., 2008; Randin et al., 2009; Meineri et al., 2012; Dobrowski 2011). Also, I investigated if species with warmer or colder continental optima than present anywhere in a focal landscape cluster at the landscape climatic boundaries, and compared the optima correlations between vascular plants and bryophytes due to their ecophysiological differences.

In the focal landscape (Fig 2 & 3), I identified 24 south-facing and 25 north-facing slopes (Fig 3). I randomly selected a 25 x 25 m plot with mature forest on each slope from 500 x 100 m grids. The plots had similar environmental heterogeneity with regards to soil type (mesic) and minimum distances to open areas, streams and vertical cliffs. Their altitudes ranged from 39 m to 385 m. I inventoried vascular plants and bryophytes and assessed their abundances within all the 49 plots. The abundance scale from 1 to 3 were: 1) sporadic (few occurrences covering < 5 %), 2) common (covering 5 – 50 %), 3) dominant (covering 50 %). I used 50 bryophytes and 96 bryophytes (see Fig 4b,c & 5b,c for examples), which occurred at >10 % of the plots, in the continental optima calculations and optima analyses. In the continental area (Fig 2), I retrieved occurrences of the focal vascular plants and bryophytes for the period 1950 to 2014 from GBIF (www.gbif.org), Swedish Artportalen (www.artportalen.se) and Norwegian Artsobservasjoner (www.artsobservasjoner.no). In the optima analyses, I only included countries, or parts of countries, with species densities of at least 1 occurrence per 1000 km2 of the 146 study species pooled. We calculated growing degree days (Fig 7b, base 5°C), minimum temperature (expressed as the yearly 5th percentile of the daily min

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temperature) and maximum temperature (expressed as the yearly 95th percentile of the daily max temperature) for a grid of the focal landscape (3066 km2) with 50 m grain size. The variables were derived by using a path model approach based on 1-year temperature measurements during June 2011 until May 2012 (Meineri et al., 2015). I also derived these variables for a c. 1000 × 1000 m grid of the continental area (4.40 × 106 km2) based on Worldclim data sets (period 1950-2000; www.worldclim.org; Hijmans et al., 2005) at a 30 seconds resolution (~1 km) (Fig 8a). Percentiles were not used

for minimum and maximum temperatures at the continental scale. I derived the landscape climatic optima for each species as their mean values of the temperature variables at their occurrences. These mean values were weighted by occurrence abundances. Their continental climatic optima were derived from MaxEnt ver 3.3.3e (www.cs.princeton.edu/~schapire/maxent; (Phillips et al., 2006)) models of the species’ continental distributions based on the three climatic variables respectively (see Fig 8b for an example). The optima were calculated from the mean values of the three

Figure 8. a) Yearly growing degree days (base 5°C, GDD5) for the continental study area calculated from Worldclim temperature data (1950-2000, www.worldclim.org). b) Example of a species distribution model derived through Maxent based on continental GDD5 for the moss Pseudotaxiphyllum elegans, which prefer rather moist conditions and have a relatively high GDD5 optima.

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temperature variables for the 1 km grid cells with 90th percentile probability of occurrence. The study species were divided into three groups (differently for each temperature variable) based on if the conditions of their continental optima was present or not in the focal landscape. Species with their continental climatic optima present in the focal landscape were denoted central species (C), species with lower continental climatic optima than present in the focal landscape were lower cluster species (LC), and species with higher continental climatic optima than present in the focal landscape were upper cluster species (UC). I used Arcmap (Version 10.1, Esri, www.esri.com) for calculations of species’ occurrence densities for countries, continental temperature variables and focal landscape climatic boundaries.

Population dynamics of a forest moss across microclimatic gradients (Paper IV) I investigated the impact of microclimatic gradients on the population dynamics of the forest moss H. umbratum (Fig 4c). I examined how population growth rate and proportion of short segments of transplants were related to evaporation, maximum temperature and snow cover duration. Also, I compared the population dynamics between northern and more southern populations.

I selected 30 grown-up Norway spruce Picea abies forests in the study area from aerial photos. I stratified their locations by including a wide range of slope aspects and degrees in order to cover a large microclimatic gradient due to variation in incoming solar radiation. In each forest, I placed a 4 ×4 meter site on mesic soil and within homogeneous slope aspect and degree. In addition, all sites were located at least 50 m from open areas and streams, 25 m from younger forest stands and 10 m from vertical cliffs in order to reduce some environmental heterogeneity. Their altitudes ranged from 158 m to 379 m and were retrieved from a digital elevation model (DEM) with a resolution of 50 x 50 meter (Lantmäteriet; the Swedish mapping, cadastral and land registration authority, www.lantmateriet.se; ArcGIS Desktop 10.1).

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I derived maximum temperature and snow cover duration for each site from ground temperature measurements with ibuttons (DS1922, (Hubbart et al., 2005)). I placed the logger underneath the moss cover inside two small, plastic zip-bags. They recorded temperature every 70 minute during the two years experimental period. I averaged the two years maximum temperatures in order to retrieve the predictor. I reduced the influence of outliers by using the 95th percentiles of maximum temperature (Ashcroft & Gollan, 2012)). I let the number of days between snow cover start and end dates represent the snow cover duration each year, and used the average of these two values per site as predictor. The snow cover duration was calculated from the ground temperature measurements in a similar way as did Lundquist & Lott (2008) and Vercauteren et al. (2013b). However, I used the 5th respectively 95th percentiles of snow cover days as the start and end dates, respectively, in order to get more robust estimates. In order to approximate humidity of each site, I measured evaporation during June until September 2013 from two narrow, c. 50 cm high plastic cylinders filled with distilled water. Water could evaporate from their bottoms through a thin filter paper similarly to the passive transpiration from a moss leaf. Each cylinder was attached to a wooden pole with its bottom 10 cm above the ground. The mean value between evaporation from the two cylinders was used as the evaporation predictor.

I gathered transplant material in late May 2012 from 6 sites (Fig 2), and attached three northern and three southern originated transplants (of about 10 cm in diameter) to each site in June 2012. The transplants were attached to rather flat ground with rubber-coated steel wires in a 2 × 0.5 m large area with 5 cm thick layer of planting peat. I counted and marked individual growing points with PVC rings (cf. Økland, 1995) during June 2012. 805 mature segments, stemming from the 2012 growing points, were measured (mm) in June 2013 and followed up in June 2014 when the next segment generation had matured. Mature segments arose from growing points, from branches following stem breaks and from apical growth. I calculated two response variables for the population dynamics analyses; population growth rate (lambda ( )) for

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segments and stable stage index for three segment size classes. I derived two values of each response variable from each site, one for the northern populations and one for the southern populations. I calculated lambda (x = 1.16; 0.13 - 2.43) for transition matrices (Økland, 1995; Rydgren & Økland, 2002), consisting of three segment size classes: S1: >0 - 15 mm, S2: >15 - 30 mm, and S3: >30 mm. From the two matrices of each site (the northern and southern population), I retrieved both lambda (the eigenvalue) and stable stage distribution (the right eigenvector). From the stable stage distribution, I retrieved the stable stage index (x = 0.70; 0 – 1) by the formula: (‘S1 proportion’ × 1) + (‘S2 proportion’ × 0.5) + (‘S3 proportion’ × 0), in order to approximate the proportion of small segments in the stable stage of each population.

Results and Discussion The overall results of this thesis suggest that microclimate influence the distribution of forest bryophytes and vascular plants by influencing their vital rates (growth, survival, reproduction) and, thus, their population growth rates. It may be possible to predict species distribution patterns at a larger, continental scale, by studying the microclimatic influence on population growth rates and distributions at a rather small, landscape scale. However, several local factors seems to influence landscape distribution patterns in a complex way, such as moisture and biotic interactions. This is important to consider, along with population differentiations, when inferring the location of e.g. microrefugia from larger distribution patterns. The results imply that forest bryophyte population growth rates and distributions at both landscape and continental scales are more affected by moisture conditions compared to forest vascular plants, for which temperature conditions seems more important.

The results of paper (I) confirmed our prediction of better performance, in terms of transplant size change, of the northern liverwort B. lycopodioides on relatively cool north-facing slopes as compared with warmer south-facing slopes. However, the two more southern distributed mosses E. angustirete and H. seligeri did not

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perform better at warmer south-facing in contrast to our hypothesis. Still, E. angustirete performed better in warm temperatures. For B. lycopopodioides the better growth at north-facing slopes could partly explain its northern distribution in Sweden. Its growth and occurrences might be favoured by the more stable moisture conditions at north-facing slopes, induced by for example colder daytime air temperatures. The relatively high performance of E. angustirete north of its range margin suggest that its distribution might not be in equilibrium with current climatic conditions (cf. Svenning & Sandel, 2013), albeit its performance over several growing seasons needs to be further examined. Its better performance in warmer conditions agree with its more southern distribution. The reason for similar growth between aspects could be that also other factors than aspect influence climatic conditions (e.g. Geiger et al., 2003). For example, altitude that influence mild maximum temperatures and might have induced variation to the aspect comparison. Also, warmer temperatures at south-facing slopes might lead to faster growth during shorter periods of time due to drought, while the about equal performance at the often colder north-facing slopes might be explained by longer periods in hydrated and growing state. From the results it is clear that there was a species-specific response to microclimatic gradients. The influence of microclimatic conditions on performance during one summer might not determine the distribution of the southern species, albeit it agreed more with the distribution of the northern species. Further, several climate-forcing factors probably influence microclimate in complex way and, thus, the performance and the distributions of bryophytes. This is corroborated by the results of paper (IV), where evaporation was most important for the performance of H. umbratum, which in turn is largely dependent on relative humidity (Papaioannou et al., 1996). Relative humidity may be favoured by for example wind-protected sites or high soil moisture, and is also influenced by temperature. From the results of paper (I) and (IV), I suggest that it would be important to investigate how the variation in relative humidity across a landscape, also in combination with

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temperature over time, influence vital rates of bryophytes in order to predict their future distributions.

The results of paper (II) contrasted the prediction that northern, marginal populations of L. vernus onset vegetative and flowering development earlier with a shorter development time than the southern populations from warmer climates, when grown in a common environment. We did not find any significant differences in the timing of development between plants from the two regions, in spite of a general population differentiation. Therefore, it seems that site- or region specific environmental conditions or genetic processes such as genetic drift was more important for population differentiation than larger scale regional effects. Our prediction for individuals within respectively region was partly confirmed. Within regions, we hypothesized that populations from colder climates and sites with higher proportion of deciduous trees develop earlier and faster than populations from warmer climates, in a common garden. We did find that plants from colder sites and higher proportion of deciduous trees in the southern region started to flower earlier, while there were no such correlations for vegetative development. In contrast, plants from colder sites in the northern range developed had earlier vegetative development, but not earlier flowering time. Within the southern region, it may be more important for individuals to flower before canopy closure, since these sites are dominated by deciduous trees. The reason would be that both light conditions and pollinator availability are probably higher before canopy closer in forests dominated by deciduous trees. The northern sites are dominated by coniferous forests so that earlier development at sites with more deciduous trees might less important. However, earlier vegetative development at colder sites might be relatively more important for population growth rates toward more harsh, marginal conditions, in order to increase net assimilation and survival (cf. Billings and Mooney 1968; Olsson and Ågren 2002; Crawford 2004). To summarize, there was earlier development in plants of populations from colder sites within both regions, when studied in the common garden. This is similar to a countergradient variation (Conover & Schultz, 1995; Conover et al., 2009), where individuals

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from colder sites are selected to develop earlier due to the shorter growing season.

Population differentiation in flowering time was mostly explained by the joint effects of start and rate of vegetative development, which were negatively correlated. This could be a reason for that reproductive and vegetative development were differently affected within the two regions. From these results, we have learnt more about how biotic and climatic factors interact to determine population differentiation in the timing of development, which might be due to local adaptation. Responses in vegetative and reproductive development of populations may differ among populations to future changes in light and temperature conditions, both across and within generations. Such knowledge could help us to better understand how vital rates and population growth rates are influenced by temporal changes in climate. The link between how local factors such as temperature and tree species composition affect population dynamics, could further elucidate which factors that determine the location for microrefugia or stepping stones (cf. Hannah et al., 2014), and may help us to explain the large variation in landscape optima in paper (III).

In paper (III), I predicted that species landscape and continental optima for bryophytes and vascular plants are positively correlated. Also, I predicted that species having their optima outside the focal landscape (either at the warmer or colder direction) would cluster at the warmest and coldest places in the landscape, respectively, and that the correlations for vascular plants should be stronger than for bryophytes due to ecophysiological differences. Our results were in line with the first prediction since there were optima correlations for all species pooled of growing degree days and maximum temperature, while there was no such correlation for minimum temperature. This suggest that we may infer species continental distributions from correlations between their smaller scale landscape distributions and local climatic factors. Species that occur at warm places in the landscape generally have more equator-ward continental distributions and vice versa. Analogously, species continental scale distributions inform us about their distributions at

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the landscape scale. Thus, climate not only regulate the distributions of sessile species at larger scales (cf. Pearson & Dawson, 2003), but also across landscapes. In contrast with our hypothesis, I did not find any clear clustering patterns since the variation in landscape optima was rather large. This suggest that also other factors, such as moisture, light and soil conditions, interact to determine species occurrences. This is not surprising when we consider the importance of both moisture conditions and temperature in papers (I) and (IV), and the effects of deciduous trees in paper (II) on individual performance. Also, vascular plants such as L. vernus in paper (II) with relatively large seeds may be dispersal limited, especially in topographic heterogeneous landscapes with abundant physical barriers (Linhart & Grant, 1996; Schiemann et al., 2000). This may enhance local adaptation, which was common for L. vernus, and suggest that dispersal limited species do not occur in all suitable climatic conditions of the landscape (Svenning & Sandel, 2013). The influence of many factors on performance may limit our ability to up- or downscale species distributions, and our ability to identify places with landscape clustering of for example species with cold continental optima. However, the landscape optima for maximum temperature was at average warmer and colder, respectively, compared to the landscape optima for central species (that have their optima within the landscape), indicating some clustering. Places in the landscape with cold maximum temperatures, e.g. on low relative elevations on north-facing slopes, may function as microrefugia during climate warming for species with colder continental optima, although our results did not clearly support such clustering. For example, the better performance of B. lycopodioides (paper (I)) at north-facing slopes indicate that such cold places might function as future microrefugia for this species. Identifying future microrefugia during climate change would favour from further studies on which environmental factors that govern landscape clustering.

The optima correlations for bryophytes were weaker and they only retained a positive correlation for maximum temperature, while vascular plants retained both correlations. This probably reflect that bryophytes often are more sensitive to water conditions, since

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desiccation of the poikilohydric bryophytes leads to shorter time for shoot growth (Goffinet et al., 2009; Proctor, 2009). Both paper (I) and paper (IV) points at the importance of water availability for the performance of bryophytes. From the results it seems likely that moisture conditions regulate the distributions, and population growth rates, of forest bryophytes not only in the focal landscape, but also across their geographical ranges in Europe. Many bryophytes are distributed toward more maritime parts of Europe with higher moisture availability compared to vascular plants (cf. Ratcliffe, 1968; Vanderpoorten & Goffinet, 2009), which was reflected in the higher continental optima for cold temperatures and lower optima for warm temperatures for bryophytes. On the other hand, the distributions of vascular plants toward more continental parts of Europe could reflect a larger influence of temperature on their distributions. They have a better water regulating capacity (Proctor, 2011), and the growing season temperatures are important for their development. They need the energy to complete flowering, seed set and to accumulate energy reserves for the winter dormancy (cf. Woodward & Williams, 1987; Prentice et al., 1992). This was partly demonstrated in paper (II), in which L. vernus plants from colder sites within two regions were adapted to earlier flowering in order to compensate for the shorter growing season.

Paper IV suggested that the population dynamics of the forest bryophyte H. umbratum is dependent on moisture availability in terms of evaporation. As I predicted, its population growth rate was higher and its segment size larger in places with low evaporation, which correspond to sites with high relative humidity. In contrast to our prediction, population growth rate and segment size were correlated neither to maximum temperature or snow cover duration. Further, there was no population differentiation in terms of population growth rate or segment size, between northern and more southern populations. Therefore, the plastic response of segment length to microclimatic conditions, was more important than genetic differentiation among the populations.

Influence of in situ measured microclimatic conditions across spatial scales on the population dynamics of bryophytes has

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rarely been studied before (cf. Økland, 1997). For H. umbratum, the variation in population growth rate imply that individuals respond to microclimatic conditions due to their ecophysiological traits, with lack of active water regulation (Goffinet et al., 2009; Proctor, 2009). This indicate that H. umbratum populations are sensitive to changes in microclimatic conditions. It could also explain the shorter segments in places with higher evaporation since water shortage lead to less growth and development constraints. This corresponds to the results in paper (I), i.e. the smaller size change of B. lycopodioides at north-facing slopes and the suggested moisture-dependent growth of E. angustirete. It seems that total evaporation during the summer is more important for the population dynamics of H. umbratum compared to shorter daytime heat events considering the lack of correlation with maximum temperature. Also, the relative humidity seems to affect the population dynamics more compared to the snow cover duration, in spite of its variation in the landscape (Vercauteren et al., 2013b) The contrasting effects of snow cover duration, with positive effects of insulation and melting water opposed by shortened growing season length, might be a reason for the lack of significant correlation with population growth rate. It could be a reason for why moderate snow cover favoured the abundances of several boreal bryophytes and vascular plants (Rasmus et al., 2011).

It is possible that a higher proportion of shorter segments explain lower population growth rates, considering longer shoots often have been related to higher population growth rate in both bryophytes and vascular plants (Eriksson, 1988; Nault & Gagnon, 1993; Økland, 1997). This also strengthens the results of paper (I) where transplant size change of B. lycopodioides and E. angustirete may be related to their population growth rates. However, shorter segment size may also be a response in order to better endure drier conditions. More smooth, dense growth forms of moss colonies are less exposed to transpiration (Proctor, 1982; Bates, 1998). If so, shorter segments could have higher fitness in drier conditions compared to longer segments and thus buffer decrease in population growth rate. On the other hand, larger shoots may better retain water (Økland, 1997, 2000). It is yet unknown if there are adaptive

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responses in segment length in bryophytes in response to temporarily drier conditions.

The lack of population differentiation among regions might be a consequence of the generally good dispersal ability of many bryophytes due to their small wind-dispersed spores (cf. Lönnell et al., 2014). There are no large physical barriers between the northern and southern regions that would prevent wind-dispersal and gene flow. Therefore, it could make more sense to generalize observed patterns across populations of more well-dispersed species, if they have not been separated by dispersal barriers (cf. II). Moreover, the elapsed time since the last glacial maximum is maybe too short for genetic diversification within perennial bryophyte species (Hedenäs, 2012).

By inferring the population growth rates of H. umbratum across microclimatic gradients from individual vital rates, I examined how microclimatic conditions affect its population dynamics and, thereby its distribution. It should be possible to map future conditions of environmental variables that affect its population growth rate, such as evaporation, correlate them to vital rates and predict its future distribution (Fig. 9). For example, by predicting occurrences in conditions where its population growth rates are positive without influence of density (cf. Diez et al., 2014; Merow et al., 2014). In addition, moisture-related variables such as evaporation probably affect the performance and distributions also of other forest bryophytes, considering the results of paper (I) and (III). Furthermore, the optima correlations of paper (III) suggest that by mapping population growth rates across local environmental conditions, via vital rates such as for H. umbratum, we may be able to infer distribution patterns at larger scales or in other parts of the species range. Similarly, we may be able to say something about the European distribution of the bryophytes in paper (I) by examining how their performance relates to vital rates.

The transplant method to determine population growth rates across environmental conditions could also be applied to other sessile organisms, in order to improve species distribution modelling. Species can be transplanted beyond and within their

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normal geographical distributions (I, IV), in order to investigate the influence of a broader range of environmental conditions on population dynamics than found within their distribution areas. For example, the conditions under which population growth rates are zero and below can be examined, i.e. the species niche limits in the absence of density effects (cf. Ehrlén & Morris, 2015). Moreover, the results of paper (I) demonstrate that after one growing season we can learn more about species performance across microclimatic gradients, even if longer transplant periods imply more climatic variation and possibilities to infer vital rates of life-stages. Distribution models based on population dynamics could be validated by using both smaller scale abundance data and larger scale presence/absence-data for the focal species (cf. III), or the other way around. Transplant experiments approximate population dynamics under relatively natural conditions, since both abiotic factors, that I assessed, and biotic factors influence the transplants, e.g. in terms of intra-specific density-dependence and influence from surrounding plants. This imply that abundances and intra-specific density-dependence could be quantified and incorporated into distribution models based on transplant experiments (Ehrlén & Morris, 2015). Thus, paper (IV) represent an important base for research on species distribution models based on population dynamics under current and future climatic conditions.

Figure 9. The distribution of a species represent the sum of its geographical occurrences, and can be predicted from the influence of environmental factors on its population dynamics. The population growth rate is set by its individual vital rates, which in turn are affected by abiotic and biotic factors. In addition, different population growth rates lead to varying abundances of species occurrences and density-dependent feedback on the vital rates. Also, dispersal (migration) and population fluctuations may affect local abundance. Adapted from Ehrlén & Morris (2015).

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Concluding remarks This thesis indicate that microclimatic gradients are important for the population dynamics of both forest bryophytes and vascular plants through their influence on vital rates and population growth rates. This suggest that we could predict species’ distributions by correlating microclimatic conditions to their population growth rates, via their vital rates. Further, population differentiation can be important to consider when studying population dynamics, but it was more important for a forest herb than for a bryophyte (II, IV), possibly due to different traits such as dispersal capacity. Moreover, local environmental conditions were more important for population differentiation of a forest herb compared to larger scale climatic gradients between regions (II). Also, species’ ecophysiological traits influence how microclimate affect their population dynamics and distributions. Bryophyte vital rates and distributions were generally more affected by moisture conditions than by temperature, as indicated by the bryophyte experiments and optima correlations (I, III, IV). Hence, they may be sensitive to altered conditions in moisture availability through for example climate change (I, IV). For vascular plants, the vital rates and distributions are probably more influenced by temperature, as suggested by a common garden study and the optima correlations (II, III). The correlations between landscape and continental optima across species further imply that we may be able to infer species’ continental distribution patterns from their landscape distributions, and vice versa (III). However, the variation in species landscape optima (III) corroborate the experimental results that several local factors influence performance, and that climate-forcing factors determine microclimate in a complex way (I, II, IV). This needs to be considered when predicting microrefugia or stepping stones from larger scale distribution patterns (cf. Hannah et al., 2014). Nonetheless, we have learnt more about which factors that are important for the distribution of species with different ecophysiological traits, by comparing optima correlations across scales for different environmental variables combined with local experiments. For a forest moss, I inferred

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population growth rates across its local distribution limits within a landscape, through the effects of evaporation on transplant vital rates. In this way, we could quantify how microclimatic factors affect population growth rates. This approach can be applied to other species as well, in order to refine our use of species distribution models. Thereby, we could in a more ecological realistic way predict species’ future, landscape distributions in response to climate change (cf. Ehrlén & Morris, 2015), and, also, infer larger scale distribution patterns (cf. III).

Acknowledgements I would like to thank my supervisor Kristoffer Hylander for valuable comments on this thesis summary.

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Svensk sammanfattning Arters utbredningar antas ofta utgöras av de platser där deras populationstillväxt är positiv. Populationstillväxt styrs av hur individernas tillväxt, reproduktion och överlevnad påverkas av olika miljöfaktorer som t.ex. klimat. Detta behöver dock studeras mer för att förtydliga bilden av hur arters utbredningar regleras. Vid ett förändrat klimat kommer de populationer som finns i utkanterna av artens utbredningsområde att vara särskilt utsatta. Klimatet vid en arts sydliga utbredningsgräns kan exempelvis bli för varmt eller torrt för att de populationen ska kunna överleva. Populationen skulle kunna överleva om den kan anpassa sig till nya klimatförhållanden, eller om det finns platser i landskapet med lite kyligare mikroklimat, så kallade mikrorefugier. Mikroklimat är det klimat som individer upplever, och utgör summan av regionala luftrörelser och lokala klimatfaktorer. På senare år har betydelsen av mikroklimat i varierad terräng för arters utbredning uppmärksammats alltmer, inte minst i skenet av klimatförändringar. Variation i mikroklimat uppkommer exempelvis genom varierande solinstrålning, höjd över havet eller avstånd till havet. I denna avhandling har jag inriktat mig på att studera hur mikroklimat inverkar på kärlväxters och mossors tillväxt, överlevnad och reproduktion och därmed deras populationstillväxt och utbredningar. Framförallt har jag studerat vilka mikroklimat som gynnar olika arter eller populationer och som begränsar arternas utbredningar.

Individer kan uppvisa olika fenotypiska karaktärer i olika miljöer, exempelvis om de växer på olika sätt eller olika mycket under olika klimatförhållanden. Detta kallas för plasticitet. Detta kanske inte räcker för populationens överlevnad i det fall klimatförändringen blir större än artens ekologiska nisch (de miljöförhållanden där populationer kan etableras och tillväxa). Population kan då överleva genom naturligt urval som anpassar dess genuppsättning, eller genom att sprida sig för att undvika det nya klimatet. I denna avhandling har jag studerat plasticitet och lokal anpassningar genom naturligt urval.

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Mossor utgörs av bladmossor, levermossor och nålfruktsmossor och härstammar från grönalger. Det var de första gröna växterna som koloniserade land för ca 400 miljoner år sedan. De är numera den näst artrikaste växtgruppen efter kärlväxter med ca 16 000 arter, och utbredda över en stor del av jorden. De är relativt artrika i boreala skogar och dominerar dess bottenskikt. Mossor är relativt lågväxta, vilket beror på att de saknar stödjande, vattentransporterande vävnad. Dessutom kan de inte reglera upptag och intag av vatten. Det gör även att de lätt torkar ut, vilket de klarar av, men som kan leda till stress och kortare period av tillväxt.

Kärlväxter har dominerat jordens flora under senaste 300 miljoner åren. Till kärlväxter räknas lummerväxter, ormbunksväxter och fröväxter. Det är den näst artrikaste organismgruppen efter skalbaggar och utgörs av omkring 300 0000 arter världen över. De bilder de vidsträckta boreala skogarna som utgör studiesystem i denna avhandling. Deras storlek och växtsätt uppvisar en stor variation. Generellt kan de bli större än mossor eftersom de har stöd av vattentransporterande vävnad, som hos buskar och träd utgörs av ved. De kan även reglera upptag och intag av vatten med bland annat rötter, vilket gör att de inte torkar ut lika lätt som mossor. De klarar dock inte av att torka ihop som exempelvis mossor och lavar.

I denna avhandling har jag studerat utbredningar och anpassningar för både kärlväxter och mossor och hur dessa organismgrupper skiljer sig åt. Jag har främst undersökt detta i boreala skogar i Ångermanland och Medelpad. Detta område har en varierad topografi som ger en stor variation i mikroklimat. Detta har bidragit till förekomster av både nordliga och sydliga kärlväxter och mossor som där befinner sig nära deras utbredningsgränser.

I artikel I transplanterade jag två sydliga bladmossor, hasselmossa Eurhynchium angustirete och stubbspretmossa Herzogiella seligeri, och en nordliga levermossa, lummermossa Barbilophozia lycopodioides, till 15 relativt varma sydsluttningar och 18 lite mer kyliga nordsluttningar i Ångermanland. Mossorna samlades in i deras huvudutbredningsområden, för de sydliga mossorna i Uppland, och de nordliga mossan i Ångermanland. Det innebär att hasselmossa transplanterades lite norr om sin

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utbredningsgräns, stubbspretmossa transplanterades nära sin utbredningsgräns och lummermossa transplanterades i sitt huvudutbredningsområde. Vi studerade därefter tillväxt, vitalitet och kapselmognad hos transplantaten under sommaren 2011. Samtidigt mätte vi temperatur och produktivitet (vegetationsklasser) samt räknade ut solinstrålning och avstånd till havet.

I artikel II samlade vi in fröer av en sydlig kärlväxt, vårärt Lathyrus vernus, från 10 nordliga utpostlokaler i Ångermanland och Medelpad och från 10 mer sydliga lokaler i Uppland, Södermanland och Östergötland. Insamlingen gjordes under 2010-2011. Vi odlade fram 375 plantor av vårärt, som vi studerade i bänkgård under våren och försommaren 2014. Vi noterade start för vegetationstillväxt, tid för vegetationsutveckling fram till blomning samt första blomningsdag. Vi räknade ut årlig maxtemperatur genom temperaturmodeller samt andel lövträd från ortofoton för alla 20 ursprungslokaler.

Under somrarna 2011 och 2012 inventerade jag förekomster och abundanser av 50 kärlväxter och 96 mossor i 25x25 meter stora rutor på 49 syd- och nordsluttningar i Ångermanland (artikel III), vilket jag kallade för landskapsnivå. Vi räknade även ut temperaturer för dessa lokaler från temperaturmodeller baserade på våra lokala temperaturmätningar. Vi modellerade fram utbredning av dessa arter på kontinental nivå (en stor del av Europa) med hjälp av förekomster i en internationell databas (GBIF) och internationella klimatmodeller (WorldClim). Vi räknade ut arternas optima för årlig maxtemperatur, mintemperatur och graddagar under tillväxtsäsongen, dvs. den temperatur där varje art förekommer mest. Detta utförde vi för både landskapsnivå och kontinental nivå.

I artikel IV transplanterade jag sydliga och mer nordliga tussar av mörk husmossa Hylocomiastrum umbratum till 30 inlandslokaler i Ångermanland och Medelpad med sluttningar i olika väderstreck, och därmed olika mikroklimat. De sydliga transplantaten samlade jag in i Västergötland i Småland, medan jag hämtade de mer nordliga transplantaten i Hälsingland och Ångermanland. Totalt 3 sydliga och 3 nordliga transplantat placerades på varje lokal. Jag mätte därefter enskilda segment

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avseende tillväxt, överlevnad och längd mellan juni 2013 och juni 2014. Mätningarna från 2013 och 2014 använde jag för att räkna ut andel små segment hos varje transplantat (population) och transplantatens (populationernas) populationstillväxter. På varje lokal mätte vi evaporation från vattenfyllda cylindrar under sommaren 2013, samt marktemperatur och snötäckets längd (baserat på temperaturmätningarna) under 2012-2014.

Resultaten från artikel I visade att den nordliga lummermossan växte bättre på nordsluttningar jämfört med sydsluttningar. Nordsluttningar i detta landskap var generellt lite kyligare och betraktas ofta som fuktigare än sydsluttningar. Jag hade därför förväntat att denna art växer bättre i lite mer kyliga och fuktiga förhållanden. Den sydliga hasselmossan växte däremot bra på både syd- och nordsluttningar. Detta var förvånande eftersom landskapet ligger norr om dess utbredningsgräns. Dess utbredning kanske inte är i jämvikt med rådande klimatförhållanden, vilket kan indikera att den kan sprida sig norrut. Alternativt så behöver vi studera transplantatens tillväxt under flera år för att få fördjupad kunskap om dess ekologiska nisch. Hasselmossan växte däremot lite bättre i varmare klimat. Det tyder på att mikroklimatet i landskapet influeras av många faktorer, t.ex. höjd över havet, och inte bara av sluttningens väderstreck. Vidare, så skulle resultaten kunna förklaras av att nordsluttningarna var mer fuktiga med längre tillväxtperiod och sydsluttningarna mer torra med kortare, men snabbare tillväxtperiod genom varmare temperaturer. Även den sydliga stubbspretmossa trivdes ungefär lika bra på nord- som sydsluttningar, vilket inte reflekterade dess geografiska utbredning.

I artikel II fann vi att det inte var någon skillnad i vegetationsstart, vegetationsutveckling och blomningsstart mellan nordliga och sydliga populationer när det studerades i en gemensam miljö. Det står i kontrast till många studier som har visat på att populationer från kallare klimat är anpassade till tidigare utveckling, för att kompensera för den kortare tillväxtsäsongen. Det finns dock andra studier som har visat på liknande resultat. Det kanske kan förklaras av att de sydliga populationerna kom från lövträdsmiljöer och de norra från barrskogsmiljöer. De sydliga populationerna skulle

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kunna vara anpassade till att utvecklas och blomma innan lövsprickningen, eftersom det då blir mindre solljus till utveckling och pollinatörer. Samtidigt skulle de nordliga populationerna också kunna vara anpassade till relativt tidig utveckling på grund av en kortare tillväxtsäsong med kallare vårar. Detta stärks av resultaten inom den sydliga regionen, där populationer från platser med större andel lövträd började blomma tidigare. Både inom den nordliga och sydliga regionen började populationer från kallare platser att utvecklas tidigare. Sammanfattningsvis så hade de lokala faktorerna inom varje region större betydelse för skillnader mellan populationer, än klimatgradienter över större skala mellan regioner.

Resultaten i artikel III visade att arters utbredningar på landskapsnivå i förhållande till klimat, dvs. klimatoptima, är korrelerade mot klimatoptima på kontinental nivå. Vi kan därmed säga något om var de finns på kontinental nivå, dvs. i Europa, genom att studera i vilka klimatförhållanden en art förekommer på landskapsnivå. Jag hittade dessa samband för maxtemperatur och graddagar, men inte för mintemperatur. Det kan ha berott på att mintemperatur och de två andra variablerna var tydligt positivt korrelerade på kontinental nivå, men inte på landskapsnivå. Det kan också bero på att årets mintemperatur infaller under vintern när många växter är skyddade av ett tjockt snötäcke i Ångermanland. Vidare fanns det en stor spridning i vilka klimat på landskapsnivå som sydliga och nordliga arter förekommer. Det innebär att det kan vara svårt att säga var sydliga arter förekommer i landskapet, eller var det kan uppstå kyligare mikrorefugier för arter vid ett kallare klimat. Detta tyder på att många faktorer, inte enbart enstaka temperaturvariabler, inverkar på arternas lokala utbredningar, till exempel fukt- och ljusförhållanden. Exempelvis hade många mossor en mer maritim utbredning i Europa än kärlväxter, vilket visar att de föredrar fuktigare miljöer med ett stabilt mikroklimat.

I artikel IV fann jag att varierande fuktförhållanden i form av evaporation inverkar på populationsdynamik hos mörk husmossa, snarare än maxtemperatur och snötäckets längd. Populationstillväxten i antal segment var högre på lokaler med låg evaporation. Detta förklaras av att låg evaporation borde gynna

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individernas fuktstatus och därmed tillväxt, produktion av nya segment och överlevnad i populationerna. Dessutom var andelen långa segment högre på platser med låg evaporation. Tid för tillväxt blir längre och fysiologisk stress från uttorkning lägre. Endast fåtal studier har gjorts där klimatförhållanden över geografiska områden har kopplats till populationsdynamik. Så vitt jag vet är detta den första studien som gör detta för lokalt uppmätta klimatförhållanden genom transplantat. Det är ett viktigt steg på vägen för att bättre förstå klimatförändringars inverkan på arters lokala utbredningsmönster.

Sammanfattningsvis indikerar denna studie att mikroklimat är viktigt för populationsdynamik av både kärlväxter och mossor. Jag fann att mikroklimat inverkar på individernas tillväxt, överlevnad och reproduktion och därmed på deras populationstillväxt. Detta visade jag genom experiment på lokal nivå, inte minst vid arters utbredningsgränser som kan ge en uppfattning om inverkan av miljöförhållanden som inte finns inom artens vanliga utbredningsområde. Sambanden mellan mikroklimat och populationsdynamik kan användas för att prediktera arternas utbredningar på landskapsnivå. Detta kan ge en förfinad bild av hur populationsdynamik och populationstillväxt är kopplat till arters utbredningar. Genom att kunskap om hur arters utbredningar i ett landskap regleras av klimat kan vi få en uppfattning om var de finns på en större kontinental nivå. Studierna tyder dock på att flera faktorer inverkar på arternas populationsdynamik, inte minst fukttillgång som generellt verkar vara viktigare för mossor i skogsmiljöer än för mer temperaturgynnade kärlväxter. Inverkan av flera faktorer kan ha bidragit till en stor spridning i arternas klimatoptima på landskapsnivå. Det försvårar möjligheten att peka ut exempelvis framtida mikrorefugier för arter under ett varmare klimat från deras större utbredningsmönster. Det kan dock vara möjligt, om vi lär oss mer om de faktorer som begränsar mikrorefugier. Genom att jämföra korrelationer mellan landskapsoptima och kontinentala optima, kombinerat med lokala experiment, har vi fått mer kunskap om vilka klimatfaktorer som styr arters utbredningar på olika skalor.

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Tack/Acknowledgements Jag vill tacka mina handledare Johan och Kristoffer för all hjälp under min doktorandperiod. Därefter vill jag tacka Elsa Fogelström för hjälp med exempelvis R, fenologi och medförfattarskap till vårärtsmanuset.

Tack till Niklas Lönnell för roliga diskussioner, introduktion till botan, hjälp och tips kring mina studier, övernattningar, flugfångster, starrkryss, mossexkursioner och den fina framsidan. Tackar Victor Johansson för roliga pratstunder, hjälp med R, botaniska upptäckter och övernattning.

Mina rumskompisar på gamla botan Lisa Fors och Alma Strandmark vill jag tacka för stöttning och roliga diskussioner. Lisa vill jag även tacka för musikaliska övningar. Tack till Malin König och Johan Dahlgren för bland annat statistikhjälp. Tack till Eric Meineri för medförfattarskap, analyshjälp (inte minst med utbredningsanalyser) och klimatdiskussioner.

Jag vill tacka Veronika Johansson och Mattias för övernattning och guidning i Solna. Tack till Caroline Greiser för nya uppslag kring mina studier och för intressant utbyte kring dessa. Tack till min företrädare Martin Schmalholz, vars förstudier i Ångermanland och Medelpad jag hade stor nytta av.

Tack till Johannes Forsberg för många skratt och outtröttlig hjälp under långa fältperioder. Tack till Anna Herrström och Jessica Oremus för hjälp med sådd, odling, fältnoteringar och trevliga pratstunder. Även tack till er andra som har hjälpt till i fält och i växthuset; Maria Enskog, Susanna Gross, Kerstin Kempe, Jelena Mocevic, Samuel Olausson, Ophélie Tran och Liselott Wilin.

Tack till alla mina andra nuvarande och före detta doktorandkollegor och postdocs som bidrog till en givande tid: Petter Andersson, Matilda Arnell, Martin Dahl, Tove von Euler, Helena Forslund, Joakim Hansen, Nils Hedberg, Maria Johansson, Bryndis Marteinsdotter, Gundula Kolb, Karin Lönngren, Ulrika Samnegård, Ellen Schagerström, Tenna Toftegaard, Thomas Verschut, Tiina Vinter samt till er nyare doktorander Pil Rasmussen och Beate Proske.

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Även tack till er äldre forskare som bidragit till en god tid och mycket lärdom, bland andra Kjell Bolmgren (för löparinspiration), Ove Eriksson och Peter Hambäck.

Jag vill tacka Leila Ahonen för hjälp med det administrativa och annat. Johan Klint vill jag tacka för datorhjälp samt musikaliska övningar. Tack till Peter och Ingela i växthuset för hjälp med såning och odling, trevliga diskussioner samt för katalpor och andra växter.

Thanks to my climate change/ecology co-authors on side-projects, in particular to the main authors Jonathan Lenoir and Saafa Wasof. I also would like to thank all the participants and teachers on the climatic change summer schools in Abisko and Finse (Stay or go – network) for a great time there.

Tack till min före detta kollegor från Länsstyrelsen för roliga stunder. Tack till Helena Persson (numera på Botaniska trädgården i Lund) för stöd, uppmuntran, goda råd och diskussioner.

Tack till Alexandre Antonelli och Thomas Appelqvist för en första visit till vetenskapen genom Guckusko-projektet för ett antal år sedan. Dessförinnan hjälpte Heidi Paltto, Björn Nordén, Thomas Appelqvist och Tomas Hallingbäck mig in i mossornas och lavarnas värld under mitt examensarbete, tack för det.

Tack till övriga vänner och släkt, framförallt till min familj Sara och Linnéa, mina bröder Per och Sebastian samt till mina föräldrar Tommy och Eva. Tack till Tommy och Olle som väckte mitt intresse för trädgård och kärlväxter.

Lastly, thanks to the strategic research programme EkoKlim that supported this work.