Questions: An advance of high-altitude treelines has been reported in response to warming climate throughout the globe. Understanding local controls on the establishment of seedlings above the treeline is essential for predicting wider patterns of treeline response from a process-based perspective. Here we investigate patterns of seedling establishment in relation to microsite conditions in an advancing treeline ecotone in the subtropics, using climate data recorded at the plant-relevant scale. We sought to determine which temperature factors were of importance, if sheltering plays an important role in seedling establishment and if the response varied with seedling age. Location:The Abies kawakamii treelines of the Central Mountain Range, Taiwan, 2800 to 3275 m a.s.l.Methods: Seedlings were monitored in plots covering a range of treeline structural forms over a period of 2 yrs. Temperatures were recorded at plant-relevant height (5 cm) above ground and depth below ground with data loggers. Microtopographic sheltering (at a 10-m scale) and surrounding vegetation were measured. The influence of the above variables on seedling number and growth was investigated using generalized linear models and linear mixed effect models, respectively.Results: Soil temperatures had more influence on seedling number than air temperature, whilst air temperature was positively associated with subsequent seedling growth. Establishment patterns were found to have a strong relationship with microtopographic sheltering, with more sheltered areas having elevated seedling numbers. Early growth may have significant implications for subsequent plant performance since smaller seedlings were more sensitive to both temperature and microtopography than larger seedlings.Conclusions: Air and soil temperatures and microtopography determine spatial patterns of seedling establishment. Our results suggest that establishment above the treeline is likely to continue as the climate warms, although advance will not be spatially uniform due to the modifying influence of topography. This variability has important implications for the persistence and extinction of alpine plant communities occurring above treeline in topographically complex systems.

Ongoing global climate change is driving widespread shifts in species distributions. Trends show frequent upwards shifts of treelines, but information on changes in montane forest below the treeline and in the tropics and subtropics is limited, despite the importance of these areas for biodiversity and ecosystem function. Patterns of species shifts in tropical and subtropical regions are likely to be more complex and individualistic than global averages suggest due to high species diversity and strong influence of competition, alongside direct climatic limitations on distributions. To address the question of how subtropical montane tree species are likely to move as climate changes, we used an extensive national forest inventory to estimate distribution shifts of 75 tree species in Taiwan by comparing the optimum elevation and range edges of adults and juveniles within species. Overall there was a significant difference in optimum elevation of adults and juveniles. Life stage mismatches suggested upward shifts in 35% of species but downward shifts of over half (56%), while 8% appeared stable. Upward elevation shifts were disproportionately common in high elevation species, whilst mid to low elevation species suggested greater variation in shift direction. Whilst previous research on mountain forest range shifts has been dominated by work addressing changes in treeline position, we show that although high elevation species shift up, below the treeline species may shift individualistically, heralding widespread changes in forest communities over coming decades. The wide variation of responses indicated is likely driven by individual species responses to interacting environmental factors such as competition, topography and anthropogenic influences across the broad range of forest types investigated. As global environmental changes continue, more detailed understanding of tree range shifts across a wide spectrum of forests will allow us to prepare for the implications of such changes for biodiversity, ecosystem function and dependent human populations.

Aim Climate change is causing shifts in the range of species worldwide. In high-altitude areas forests are often observed to be shifting their upper limits to higher altitudes in response to warming temperatures. Although this phenomenon is well described, the possible consequences of this for the wider forest community have not been fully considered. In this study, we used epiphytic macro-lichens to investigate the impacts of tree line advance on associated organisms. We hypothesized that the rate of forest advance should influence the species richness and composition of associated communities.Location The Hehuanshan area, Central Mountain Range, Taiwan (24°08-09Methods Lichen communities were sampled on trees occurring at 33 tree line sites displaying varying degrees of advance. Habitat variables were recorded. ANOVA, GLMM, nestedness analysis and indicator species analysis were used to investigate patterns of species richness and community composition and their association with tree line advance and habitat variables.Results Species richness was lower in tree lines exhibiting rapid advance; reductions were related to tree size (considered here as a proxy for age) and the distance over which advance had occurred. Community composition varied with tree line form and forest position. Only a subset of species found in slowly advancing or stable forest edges occurred in rapidly advancing tree lines.Main conclusions Differential migration rates between co-occurring species and differences in habitat structure associated with tree line advance can result in community change, but this depends, amongst other factors, on the speed of tree line advance. The importance of advance rate in determining the response suggests that reductions in species richness at tree line could be transitory. However, this will depend on whether advance continues, and on the changes in habitat associated with advance. Given the complexity of tree line behaviour, the findings that we report represent an essential step in understanding community responses to climate change. This understanding is of importance for biodiversity and conservation, especially given the high rate of endemism reported for this and other alpine regions.

Aim: Plant functional traits are broadly used to quantify and predict impacts of climate change on vegetation. However, high intraspecific trait variation can bias mean values when few measurements are available. Here, we determine the extent of individual leaf trait variation and covariation across a highly heterogeneous environmental gradient for a widely distributed subtropical pine. We demonstrate the implications of trait variation for characterising species by assessing data availability and variability across the Pinus genus.Location: Central Mountain Range, Taiwan.Taxon: Pinus taiwanensis Hayata (Pinaceae). Methods:We measured eight functional traits suggested to reflect plant strategies: needle length, area, thickness, dry and fresh mass, stomatal row density (SD), leaf dry matter content (LDMC) and specific leaf area (SLA). We examined trait variation in response to climatic and physiographic factors across an elevational gradient of 495-3106 m a.s.l. using linear mixed effects models (LMMs). Intraspecific trait covariation was explored using principal component analyses (PCAs) and LMMs. Descriptive statistics were calculated for Pinus records in the global TRY plant trait database.Results: Intraspecific variability among traits was high (CV 20%-44%) and predictable with elevation (generally p < 0.05, with declining needle size and LDMC with elevation and increasing SD). However, 41%-92% of variance was un-explained by topography. Sixty-five percent of variation was explained by two trait covariation axes, with predictable changes with elevation (p < 0.001). Pinus data availability in TRY was low.Across traits, only 12.5%-53% of species had sufficient sample sizes for intraspecific analyses. Main conclusions:We show substantial trait variation for a single species, here likely driven by temperature differences and additional biotic and abiotic drivers across the elevational range. Improved understanding of the extent and implications of intraspecific variability is necessary for reliable quantifications and predictions of the impacts of environmental change, especially in understudied, hyper-diverse ecosystems such as tropical forests.

No abstract