Abstract Air and soil temperatures, snow cover, serious snow load damage to coniferous trees, wind, topography and edaphic factors on the fells situated between 67°N and 68°N in Finnish Lapland are described and their influence on the location of the actual timberline and treeline is discussed. In addition the relation between annual climate conditions and pollen deposition in the timberline ecotone is analysed and the results of seedling density monitoring in the same environment are presented. The potential for the actual timberline and treeline to advance to a higher elevation is also discussed. The field studies were carried out on the fells of Aakenustunturi, Yllästunturi and Pyhätunturi. The average altitude of the actual timberline varies from 370 metres to 402 metres a.s.l. The actual timberline is hardly ever composed of a single tree species but featured alternating occurrences of Norway spruce (Picea abies), Scots pine (Pinus sylvestris) and mountain birch (Betula pubescens ssp. czerpanovii). The mean tetratherms on the southern and northern slopes (+10.3°C and +10.1°C, respectively), the mean maximum tetratherm on the southern slope (+15.1°C) and the corresponding measures for the treeline (460 m a.s.l), the minimum tetratherm (+6.3°C), mean July temperature (+12.6°C), biotemperature (+3.3°C) and minimum effective temperature sum (455 d.d.), coincide closest with the results of earlier studies. The maximum altitudes of the actual timberline are dictated by many climatic factors on southern and western slopes with a gentle inclination, and the forest cover gradually becomes thinner, in which case the actual timberline does not form any easily distinguishable line. The lowest altitudes of the actual timberline are the results of an extremely high proportion of block fields, slope steepness and snow patches on the northern and eastern slopes. On the precipitous and rocky slopes trees have difficulties in taking root and in obtaining nutrients and water, while as a consequence of snow patches the growing season may be too short for tree growth at all, and if trees exist there they are suffering from low soil temperature and parasitic snow fungi. Serious snow load damage to trees evidently hampers any advance in the actual timberline, as do avalanches and mires. The location of the treeline is the result of a combination of a great number of unfavourable conditions for tree regeneration, seedling establishment and tree growth, such as inadequate snow protection, extreme soil temperatures, almost total destruction of trees by the snow load, wind pressure, an often inadequate effective temperature sum and length of the growing season, night frost in early summer, and poor, dry soil suffering from excessive evaporation. Actual timberline responses to predicted climate warming will differ greatly from site to site in relation to the local topography, edaphic features and associated ecological limitations. Any advance in the treeline to a higher elevation is likely to be slower and at least less certain than that in the actual timberline. In addition, advances in the actual timberline and treeline may even be prevented by phenomena occurring along with climate change. A potential key factor in this is serious snow load damage to the trees.

Against the background of past, recent and future climate change, the present thesis addresses elevational shifts of alpine treelines in the Swedish Scandes. By definition, treeline refers to the elevation (m a.s.l.) at a specific site of the upper trees of a specific tree species, at least 2 m tall. Based on historical records, the first part of the thesis reports and analyzes the magnitude of treeline displacements for the main trees species (Betula pubescens ssp. czerepanovii, Picea abies and Pinus sylvestris) since the early 20th century. The study covered a large and heterogeneous region and more than 100 sites. Concurrent with temperature rise by c. 1.4 °C over the past century, maximum treeline advances of all species amount to about 200 m. That is virtually what should be predicted from the recorded temperature change over the same period of time. Thus, it appears that under ideal conditions, treelines respond in close equilibrium with air temperature evolution. However, over most parts of the landscape, conditions are not that ideal and treeline upshifts have therefore been much smaller. The main reason for that discrepancy was found to be topoclimatic constraints, i.e. the combined action of geomorphology, wind, snow distribution, soil depth, etc., which over large parts of the alpine landscape preclude treelines to reach their potential thermal limit. Recorded treeline advance by maximum 200 m or so over the past century emerges as a truly anomalous event in late Holocene vegetation history. The second part of the thesis is focused more on long-term changes of treelines and one specific and prevalent mechanism of treeline change. The first part of the thesis revealed that for Picea and Betula, treeline shift was accomplished largely by phenotypic transformation of old-established stunted and prostrate individuals (krummholz) growing high above the treeline. In obvious response to climate warming over the past century, such individuals have transformed into erect tree form, whereby the treeline (as defined here) has risen. As a means for deeper understanding of this mode of positional treeline change, extant clonal spruces, growing around the treeline, were radiocarbon dated from megafossil remains preserved in the soil underneath their canopies. It turned out that Picea abies in particular may attain almost eternal life due to its capability for vegetative reproduction and phenotypic plasticity. Some living clones were in fact inferred to have existed already 9500 years ago, and have thus persisted at the same spot throughout almost the entire Holocene. This contrasts with other tree species, which have left no living relicts from the early Holocene, when they actually grew equally high as the spruce. Thereafter they retracted by more than 300 m in elevation supporting that also on that temporal scale, treelines are highly responsive to climate change. The early appearance of Picea in the Scandes, suggests that Picea “hibernated” the last glacial phase much closer to Scandinavia than earlier thought. It has also immigrated to northern Sweden much earlier than the old-established wisdom. The experiences gained in this thesis should constitute essential components of any model striving to the project landscape ecological consequences of possible future climate shifts.

Altitudinal treeline advance represents a sensitive and well-studied example of species response to climate warming. Although a great deal of work has been conducted globally, few studies have considered subtropical alpine treelines and little is known about their structure and function. This research aims to investigate the response of high altitude forests in Taiwan to climate variation by characterising treeline advance in the area, exploring the mechanisms driving the advance, and considering the consequences of advance for the wider community. The thesis consists of a general introduction to the topic followed by a series of papers, exploring: (1) Possible consequences of treeline shifts for biodiversity and ecosystem function. (2) The advance of the Abies kawakamii treeline through aerial photograph analysis. (3) The changes in growth rate of Abies kawakamii at treeline and the influence of altitude and temperature on growth. (4) Regeneration patterns at treeline and the importance of microclimate and topographic sheltering. (5) Consequences of the range shift for the wider forest community. The work is then concluded with a general discussion and synthesis. The main aims of this work are therefore to characterise and understand the pattern and pace of treeline advance and forest structural change throughout the Central Mountain Range of Taiwan. Treeline advance is characterised through the study of repeat aerial photographs and the mechanisms behind the observed shift are explored through the study of two key responses associated with forest advance: tree growth at treeline and seedling establishment beyond treeline. The consequences of treeline advance for the wider subalpine community are investigated through the study of epiphytic lichen communities at treeline sites. This investigation of an understudied region will allow for improved understanding of treeline response at a global scale.