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Studies of elevation clines in diversity and composition of ecological communities date back to the origins of biogeography. A modern resurgence of interests in these elevational clines is likely to contribute important insights for developing a more general theory of species diversity. In order to gain a more comprehensive understanding of geographical clines in diversity, the research programme for montane biogeography should include statistically rigorous tests of apparent patterns, comparisons of patterns among regions and taxonomic or ecological groups of species, and analyses of clines in environmental variables concurrent with biogeographical surveys. The conceptual framework for this research programme should be based on the assumption that elevational gradients in species diversity result from a combination of ecological and evolutionary processes, rather than the presumed independent effects of one overriding force. Given that montane ecosystems are hot spots of biological diversity, an expanded and integrated programme for biogeographic surveys in montane regions should provide valuable insights for conservation biologists.

Aim Species are responding to climate warming by shifting their distributions toward historically cooler regions, but the degree to which expansions at cool range limits are balanced by contractions at warm limits is unknown. We synthesized published data documenting shifts at species’ warm versus cool range limits along elevational gradients to (a) test classic ecological theory that predicts temperature more directly influences species’ cool range limits than their warm range limits, and (b) determine how warming‐associated shifts have changed the extent and area of species’ elevational distributions. Location Global. Time period 1802–2012. Major taxa studied Vascular plants, endotherms, ectotherms. Methods We compiled a dataset of 975 species from 32 elevational gradients for which range shifts have been measured at both warm and cool range limits. We compared the magnitude and variance of shifts at species’ warm versus cool limits, and quantified how range shifts have impacted species’ elevational extents and areas. Results On average species have shifted upslope associated with temperature increases at both warm and cool limits (warm limit: 92 ± 455 m/C; cool limit: 131 ± 465 m/C; overall mean ± SD). There was no systematic difference in the magnitude or variance of shifts at warm versus cool limits and thus no indication that cool limits are more directly controlled by temperature. Species’ elevational extents and available area significantly decreased for mountaintop species. Main conclusions Our results do not support the long‐standing hypothesis that cool limits are more sensitive or responsive to temperature. We find that, across the globe, mountaintop species’ ranges are significantly shrinking as they shift upslope, supporting predictions that high elevation species are especially vulnerable to temperature increases. Our synthesis highlights the extreme variation in species’ distributional responses to warming, which may indicate that biotic interactions play a more prominent role in setting range limits than previously thought.

• Efforts to develop mechanistic tree growth models are hindered by the uncertainty of whether and when tree growth responses to environmental factors are driven by carbon assimilation or by biophysical limitations of wood formation. • In this study, we used multiannual weekly wood-formation monitoring of two conifer species (Larix decidua and Picea abies) along a 900m elevational gradient in the Swiss Alps to assess the biophysical effect of temperature and water potential on wood formation. To this end, we developed a model that simulates the effect of water potential on turgor-driven cambial division, modulated by the effect of temperature on enzymatic activity. • The model reproduced the observed phenology of tracheid production, as well as intra- and interannual tracheid production dynamics of both species along the elevational gradient, although interannual model performance was lower. We found that temperature alone explains the onset of tracheid production, yet water potential appears necessary to predict the ending and the total amount of tracheids produced annually. • We conclude that intra-annual cambial activity is strongly constrained by both temperature and water potential at all elevations, independently of carbon assimilation. At the interannual scale, biophysical constraints likely interact with other factors.

Elucidating the diversity patterns and elevational distribution of medicinal plants is essential for biodiversity conservation and sustainable utilization in montane ecosystems. However, the variation in diversity across forest vertical layers and its responses to environmental gradients in mid-subtropical regions remain poorly understood. We surveyed medicinal plants in 93 plots across five elevational gradients (450-1800 m) in Meihua Mountain National Nature Reserve, China. Species composition and community structure in the tree, shrub, and herb layers were analyzed using importance values, -diversity indices, and β-diversity metrics. Principal coordinate analysis (PCoA) and correlation analysis were performed to examine diversity patterns and identify their environmental drivers, respectively. A total of 503 medicinal plant species (267 genera, 98 families) were recorded. Species richness followed the order shrub > herb > tree layer. Community composition showed clear elevational replacement. -diversity declined overall with elevation, with significant unimodal patterns in shrub and herb layers but no clear trend in the tree layer. Species turnover was highest in the shrub layer and lowest in the tree layer, peaking at low to mid elevations. Elevation exerted stronger effects on -diversity in shrub and herb layers than in the tree layer. Canopy closure was negatively correlated with shrub-layer -diversity, whereas longitude and slope were positively associated with tree- and herb-layer diversity, respectively. This study demonstrate that medicinal plant diversity in mid-subtropical mountains is jointly shaped by vertical stratification and environmental filtering. The higher sensitivity of shrub and herb layers highlights their key role in maintaining biodiversity, whereas tree-layer stability reflects greater resistance to environmental variation. Canopy structure regulates understory diversity, with additional layer-specific effects from topographic and spatial factors. These results provide new insights into diversity maintenance mechanisms and offer guidance for conservation and forest management of medicinal plant resources.

Plants have the potential to adjust the configuration of their hydraulic system to maintain its function across spatial and temporal gradients. Species with wide environmental niches provide an ideal framework to assess intra-specific xylem adjustments to contrasting climates. We aimed at assessing how xylem structure in the widespread species Nothofagus pumilio varies across combined gradients of temperature and moisture, and to which extent within-individual variation contributes to population responses across environmental gradients. We characterized xylem configuration in branches of N. pumilio trees at five sites across an 18° latitudinal gradient in the Chilean Andes, sampling at four elevations per site. We measured vessel area, vessel density and the degree of vessel grouping. We also obtained vessel diameter distributions and estimated the xylem-specific hydraulic conductivity. Xylem traits were studied in the last five growth rings to account for within-individual variation. Xylem traits responded to changes in temperature and moisture, but also to their combination. Reductions in vessel diameter and increases in vessel density suggested increased safety levels with lower temperatures at higher elevation. Vessel grouping also increased under cold and dry conditions, but changes in vessel diameter distributions across the elevational gradient were site-specific. Interestingly, the estimated xylem-specific hydraulic conductivity remained constant across elevation and latitude, and an overwhelming proportion of the variance of xylem traits was due to within-individual responses to year-to-year climatic fluctuations, rather than to site conditions. Despite conspicuous adjustments, xylem traits were coordinated to maintain the hydraulic function constant under a wide range of conditions. This, combined with the within-individual capacity for responding to year-to-year climatic variations, may have the potential to increase forest resilience against future environmental changes.

Mountains are pivotal to maintaining habitat heterogeneity, global biodiversity, ecosystem functions and services to humans. They have provided classic model natural systems for plant and animal diversity gradient studies for over 250 years. In the recent decade, the exploration of microorganisms on mountainsides has also achieved substantial progress. Here, we review the literature on microbial diversity across taxonomic groups and ecosystem types on global mountains. Microbial community shows climatic zonation with orderly successions along elevational gradients, which are largely consistent with traditional climatic hypotheses. However, elevational patterns are complicated for species richness without general rules in terrestrial and aquatic environments and are driven mainly by deterministic processes caused by abiotic and biotic factors. We see a major shift from documenting patterns of biodiversity towards identifying the mechanisms that shape microbial biogeographical patterns and how these patterns vary under global change by the inclusion of novel ecological theories, frameworks and approaches. We thus propose key questions and cutting-edge perspectives to advance future research in mountain microbial biogeography by focusing on biodiversity hypotheses, incorporating meta-ecosystem framework and novel key drivers, adapting recently developed approaches in trait-based ecology and manipulative field experiments, disentangling biodiversity–ecosystem functioning relationships and finally modelling and predicting their global change responses.

1 It is widely accepted that tropical lowland rain forest holds the greatest diversity of organisms, and it is often implied that this general pattern is also true for virtually all individual higher-level taxa. Standardized elevational transect surveys of non-flying small mammals (Insectivora and Rodentia) on geologically old, species-rich islands in the Philippines consistently show maximum diversity and relative abundance in upper montane/lower mossy forest at 1500-2200 m, often exceeding lowland species richness and relative abundance by a factor of three or more. 2 On mountains where maximum elevation exceeds 2000 m, there is a decline in species richness above about 1500-2000 m, yielding a curvilinear pattern of species richness along the elevational gradient. The peak in species richness occurs at the area of transition from montane to mossy forest, which is also the point at which rainfall probably peaks. In parallel with species richness, relative abundance of small mammals in the Philippines also increases from the lowlands to 1500-2200 m, increasing by a factor from two to 10. 3 Twelve hypotheses concerning patterns of diversity along elevational gradients, plus the null hypothesis, are evaluated. The null hypothesis of no variation and the hypotheses that maximum diversity is in the lowlands, that diversity is highest in areas of least perturbation, and that diversity increases with increasing area, are rejected. There is weak or ambiguous support for the hypotheses that diversity is greatest in areas of community interdigitation, that diversity is highest in the area of highest productivity, that diversity is correlated with habitat complexity, and that diversity is correlated with habitat diversity. There is strongest support for the hypotheses that diversity is correlated with annual rainfall, with total abundance of individuals in the community, with food resource diversity, with areas of reduced competition from other organisms, and with areas characterized by high rates of speciation. 4 Causality is difficult to evaluate because many hypotheses make non-exclusive predictions, they probably represent non-independent aspects of causal factors (in other words, there is much interaction among the processes highlighted by the various hypotheses), and they represent the range from proximate to ultimate and from descriptive to causal. All the hypotheses probably represent phenomena that exist in nature, but few (or none) represent phenomena found in all taxa. The primary challenge in the future will not be simply to accept or reject individual hypotheses, but rather to determine the circumstances under which the various causal factors are most important, how they interact, and how they can be combined into a more comprehensive and general multi-factorial model.

Litter decomposition in mountainous forest ecosystems is an essential process that affects carbon and nutrient cycling. However, the contribution of litter decomposition to terrestrial ecosystems is difficult to estimate accurately because of the limited comparability of different studies and limited data on local microclimatic and non‐climatic factors. Here, we designed a coordinated experiment within subtropical and tropical forests across ten mountains to evaluate variation in litter decomposition rates and stabilization. We tested whether elevations, soil microclimate, soil physiochemistry, tree species diversity, and microhabitat affect decomposition rates and stabilization by using the Tea bag index as a standardized protocol. We found that the associations of decomposition rates and stabilization with elevation and each environmental factor varied between mountains. Elevation significantly affected decomposition rates and stabilization in the western mountains, where soil microclimate also played a dominant role due to relatively cold environments. Across all mountains, decomposition rates decreased while stabilization increased with increasing elevation. In terms of microclimate, decomposition rates increased with increasing soil temperature and temperature variation during the growing season, whereas stabilization decreased with increasing soil temperature and moisture variation. In terms of non‐climatic factors, decomposition rates increased with increasing tree species diversity, whereas stabilization decreased with soil pH and slope. Our findings enhance the general understanding of how different factors control forest litter decomposition, highlighting the dominant role of soil microclimate in controlling carbon and nutrient cycling in cold environments and high elevations.

Shifts in soil microbial communities over altitudinal gradients and the driving factors are poorly studied. Their elucidation is indispensable to gain a comprehensive understanding of the response of ecosystems to global climate change. Here, we investigated soil archaeal, bacterial, and fungal communities at four Alpine forest sites representing a climosequence, over an altitudinal gradient from 545 to 2000 m above sea level (asl), regarding abundance and diversity by using qPCR and Illumina sequencing, respectively. Archaeal community was dominated by Thaumarchaeota, and no significant shifts were detected in abundance or community composition with altitude. The relative bacterial abundance increased at higher altitudes, which was related to increasing levels of soil organic matter and nutrients with altitude. Shifts in bacterial richness and diversity as well as community structure (comprised basically of Proteobacteria, Acidobacteria, Actinobacteria, and Bacteroidetes) significantly correlated with several environmental and soil chemical factors, especially soil pH. The site at the lowest altitude harbored the highest bacterial richness and diversity, although richness/diversity community properties did not show a monotonic decrease along the gradient. The relative size of fungal community also increased with altitude and its composition comprised Ascomycota, Basidiomycota, and Zygomycota. Changes in fungal richness/diversity and community structure were mainly governed by pH and C/N, respectively. The variation of the predominant bacterial and fungal classes over the altitudinal gradient was the result of the environmental and soil chemical factors prevailing at each site.