Globally, treeline ecotones vary from abrupt lines to extended zones of increasingly small, stunted and/or dispersed trees. These spatial patterns contain information about the processes that control treeline dynamics. Describing these patterns consistently along ecologically meaningful dimensions is needed for generalizing hypotheses and knowledge about controlling processes and expected treeline shifts globally. However, existing spatial categorizations of treelines are very loosely defined, leading to ambiguities in their use and interpretation. To help better understand treeline‐forming processes, we present a new framework for describing alpine treeline ecotones, focusing on hillside‐scale patterns, using pattern dimensions with distinct indicative values: 1) the spatial pattern in the x‐y plane: a) decline in tree cover, and b) change in the level of clustering. Variation along these dimensions results in more or less ‘discrete' or ‘diffuse' treelines with or without islands. These patterns mainly indicate demographic processes: establishment and mortality. 2) Changes in tree stature: a) decline in tree height, and b) change in tree shape. Variation along these dimensions results in more or less ‘abrupt' or ‘gradual' treelines with or without the formation of environmental krummholz. These patterns mainly indicate growth and dieback processes. Additionally, tree population structure can help distinguish alternative hypotheses about pattern formation, while analysing the functional composition of the ecotonal vegetation is essential to understand community‐level processes, controlled by species‐specific demographic processes. Our graphical representation of this framework can be used to place any treeline pattern in the proposed multi‐dimensional space to guide hypotheses on underlying processes and associated dynamics. To quantify the dimensions and facilitate comparative research, we advocate a joint effort in gathering and analysing spatial patterns from treelines globally. The improved recognition of treeline patterns should allow more effective comparative research and monitoring and advance our understanding of treeline‐forming processes and vegetation dynamics in response to climate warming.
What would current ecosystems be like without the impact of mankind? This question, which is critical for ecosystem management, has long remained unanswered due to a lack of present-day data from truly undisturbed ecosystems. Using mountaineering techniques, we accessed pristine relict ecosystems in the Peruvian Andes to provide this baseline data and compared it with the surrounding accessible and disturbed landscape. We show that natural ecosystems and human impact in the high Andes are radically different from preconceived ideas. Vegetation of these ‘lost worlds’ was dominated by plant species previously unknown to science that have become extinct in nearby human-affected ecosystems. Furthermore, natural vegetation had greater plant biomass with potentially as much as ten times more forest, but lower plant diversity. Contrary to our expectations, soils showed relatively little degradation when compared within a vegetation type, but differed mainly between forest and grassland ecosystems. At the landscape level, a presumed large-scale forest reduction resulted in a nowadays more acidic soilscape with higher carbon storage, partly ameliorating carbon loss through deforestation. Human impact in the high Andes, thus, had mixed effects on biodiversity, while soils and carbon stocks would have been mainly indirectly affected through a suggested large-scale vegetation change.
We studied tree height in stands of high-Andean Polylepis forests in two cordilleras near Cuzco (Peru) with respect to variations in human impact and climatic conditions, and compared air and soil temperatures between qualitatively defined dry and humid slopes. We studied 46 forest plots of 100 m2 of five Polylepis species at 3560-4680 m. We measured diameter at breast height (dbh) and tree height in the stands (1229 trees in total), as well as air and soil temperatures in a subset of plots. The data was analysed combining plots of given species from different sites at the same elevation (±100 m). There was no elevational decrease of mean maximum tree height across the entire data set. On humid slopes, tree height decreased continuously with elevation, whereas on dry slopes it peaked at middle elevations. With mean maximum tree heights of 9 m at 4530 m on the humid slopes and of 13 m at 4650 m on the dry slopes, we here document the tallest high-elevation forests found so far worldwide. These highest stands grow under cold mean growing season air temperatures (3.6 °C and 3.8 °C on humid vs. dry slopes) and mean growing season soil temperatures (5.1 °C vs. 4.6 °C). Mean annual air and soil temperature both decreased with elevation. Dry slopes had higher mean and maximum growing season air temperatures than humid slopes. Mean annual soil temperatures did not significantly differ and mean annual air temperatures only slightly differed between slopes. However, maximum air temperatures differed on average by 6.6 K between dry and humid slopes. This suggests that the differences in tree height between the two slopes are most likely due to differences in solar radiation as reflected by maximum air temperatures. Our study furthermore provides evidence that alpine Polylepis treelines grow under lower temperature conditions than global high-elevation treelines on average, suggesting that Polylepis species may have evolved special physiological adaptations to low temperatures.
Plant functional traits can be genetically determined or phenotypically plastic. We assessed the degree of genetic determinism in the functional traits of Andean Polylepis tree species among 14 important traits that enable the species to withstand cold and dry conditions. We conducted a common garden experiment and related the species-specific means of the functional traits to the variables of climatic niche optima of the species (mean annual temperature and annual precipitation), deducing that if the interspecific variation in the functional trait is related to the species climatic niche optima according to the theoretically-expected pattern of climate-trait relationship, the variation of the trait must be genetically determined. In general, the traits were related either to species temperature or precipitation optima. For example, leaf size, maximum photosynthesis rate and root tip abundance were related to temperature, whereas light compensation and light saturation points were related to precipitation. Only leaf size showed a significant phylogenetic signal, indicating that most of the manifested climate–trait relationships are not caused purely by phylogeny, but are mainly a result of species specialisation along an environmental gradient. However, in many cases the relationships were rather weak. This suggests that important functional traits of Polylepis species involve both genetic and phenotypic components aiming to maximise the overall fitness of the species at high elevations.