Aim: Higher-elevation areas on islands and continental mountains tend to be separated by longer distances, predicting higher endemism at higher elevations; our study is the first to test the generality of the predicted pattern. We also compare it empirically with contrasting expectations from hypotheses invoking higher speciation with area, temperature and species richness. Location: 32 insular and 18 continental elevational gradients from around the world. Methods: We compiled entire floras with elevation-specific occurrence information, and calculated the proportion of native species that are endemic ('percent endemism') in 100 m bands, for each of the 50 elevational gradients. Using generalized linear models, we tested the relationships between percent endemism and elevation, isolation, temperature, area and species richness. Results: Percent endemism consistently increased monotonically with elevation, globally. This was independent of richness-elevation relationships, which had varying shapes but decreased with elevation at high elevations. The endemism-elevation relationships were consistent with isolationrelated predictions, but inconsistent with hypotheses related to area, richness and temperature. Main conclusions: Higher per-species speciation rates caused by increasing isolation with elevation are the most plausible and parsimonious explanation for the globally consistent pattern of higher endemism at higher elevations that we identify. We suggest that topography-driven isolation increases speciation rates in mountainous areas, across all elevations, and increasingly towards the equator. If so, it represents a mechanism that may contribute to generating latitudinal diversity gradients in a way that is consistent with both present-day and palaeontological evidence.

Summary Climate and topography are among the most fundamental drivers of plant diversity. Here, we assessed the importance of climate and topography in explaining diversity patterns of species richness, endemic richness and endemicity on the landscape scale of an oceanic island and evaluated the independent contribution of climatic and topographic variables to spatial diversity patterns. We constructed a presence/absence matrix of perennial endemic and native vascular plant species (including subspecies) in 890 plots on the environmentally very heterogeneous island of La Palma, Canary Islands. Species richness, endemic richness and endemicity were recorded, interpolated and related to climate (i.e. variables describing temperature, precipitation, variability and climatic rarity) and topography (i.e. topographic complexity, solar radiation, geologic age, slope and aspect). We used multimodel inference, spatial autoregressive models, variance partitioning and linear regression kriging as statistical methods. Species richness is best explained by both climatic and topographic variables. Topographic variables (esp. topographic complexity and solar radiation) explain endemic richness, and climatic variables (esp. elevation/temperature and rainfall seasonality) explain endemicity. Spatial patterns of species richness, endemic richness and endemicity were in part geographically decoupled from each other. Synthesis. We identified several topography‐dependent processes ranging from evolutionary processes (micro‐refugia, in situ speciation, pre‐adaptation to rupicolous conditions, dispersal limitations) to human‐induced influences (introduced herbivores, fire, land use) that possibly shape the endemic richness pattern of La Palma. In contrast, climate mainly drives endemicity, which is connected to ecological speciation and specialization to local conditions. We highlight the importance of incorporating climatic variability into future studies of plant species diversity and endemism. The spatial incongruence in hot spots of species richness, endemic richness and endemicity emphasizes the need for an integrated conservation approach acknowledging different diversity measures to protect the complete spectrum of diversity. High‐elevation islands such as La Palma are highly suitable to study drivers of diversity and endemism, as they offer environmental gradients of continental magnitude on the landscape scale of a single climatic mini‐continent and a large array of in situ‐speciated endemics.

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Despite islands contributing only 6.7% of land surface area, they harbor ~20% of the Earth’s biodiversity, but unfortunately also ~50% of the threatened species and 75% of the known extinctions since the European expansion around the globe. Due to their geological and geographic history and characteristics, islands act simultaneously as cradles of evolutionary diversity and museums of formerly widespread lineages—elements that permit islands to achieve an outstanding endemicity. Nevertheless, the majority of these endemic species are inherently vulnerable due to genetic and demographic factors linked with the way islands are colonized. Here, we stress the great variation of islands in their physical geography (area, isolation, altitude, latitude) and history (age, human colonization, human density). We provide examples of some of the most species rich and iconic insular radiations. Next, we analyze the natural vulnerability of the insular biota, linked to genetic and demographic factors as a result of founder events as well as the typically small population sizes of many island species. We note that, whereas evolution toward island syndromes (including size shifts, derived insular woodiness, altered dispersal ability, loss of defense traits, reduction in clutch size) might have improved the ability of species to thrive under natural conditions on islands, it has simultaneously made island biota disproportionately vulnerable to anthropogenic pressures such as habitat loss, overexploitation, invasive species, and climate change. This has led to the documented extinction of at least 800 insular species in the past 500 years, in addition to the many that had already gone extinct following the arrival of first human colonists on islands in prehistoric times. Finally, we summarize current scientific knowledge on the ongoing biodiversity loss on islands worldwide and express our serious concern that the current trajectory will continue to decimate the unique and irreplaceable natural heritage of the world’s islands. We conclude that drastic actions are urgently needed to bend the curve of the alarming rates of island biodiversity loss.

Protected areas (PA) are refugia of biodiversity. However, anthropogenic climate change induces a redistribution of life on Earth that affects the effectiveness of PAs. When species are forced to migrate from protected to unprotected areas to track suitable climate, they often face degraded habitats in human-dominated landscapes and a higher extinction threat. Here, we assess how climate conditions are expected to shift within the world’s terrestrial PAs (n = 137,432). PAs in the temperate and northern high-latitude biomes are predicted to obtain especially high area proportions of climate conditions that are novel within the PA network at the local, regional and global scale by the end of this century. These PAs are predominantly small, at low elevation, with low environmental heterogeneity, high human pressure, and low biotic uniqueness. Our results guide adaptation measures towards PAs that are strongly affected by climate change, and of low adaption capacity and high conservation value.

Treeline research has strongly focused on mountain systems on the mainland. However, island treelines offer the opportunity to contribute to the global framework on treeline elevation due to their island‐specific attributes such as isolation, small area, low species richness and relative youth. We hypothesize that, similar to the mainland, latitude‐driven temperature variation is the most important determinant of island treeline elevation on a global scale. To test this hypothesis, we compared mainland with island treeline elevations. Then we focused 1) on the global effects of latitude, 2) on the regional effects of island type (continental vs oceanic islands) and 3) the local effects of several specific island characteristics (age, area, maximum island elevation, isolation and plant species richness). We collected a global dataset of islands (n = 86) by applying a stratified design using GoogleEarth and the Global Island Database. For each island we extracted data on latitude and local characteristics. Treeline elevation decreased from the mainland through continental to oceanic islands. Island treeline elevation followed a hump‐shaped latitudinal distribution, which is fundamentally different from the mainland double‐hump. Higher maximum island elevation generated higher treeline elevation and was found the best single predictor of island treeline elevation, even better than latitude. Lower island treeline elevation may be the result of a low mass elevation effect (MEE) influencing island climates and an increasingly impoverished species pool but also trade wind inversion‐associated aridity. The maximum island elevation effect possibly results from an increasing mass elevation effect (MEE) with increasing island elevation but also range shifts during climatic fluctuations and the summit syndrome (i.e. high wind speeds and poor soils in peak regions). Investigating islands in treeline research has enabled disentangling the global effect of latitude from regional and local effects and, at least for islands, a comprehensive quantification of the MEE.

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Aim Mountains and islands are both well known for their high endemism. To explain this similarity, parallels have been drawn between the insularity of “true islands” (land surrounded by water) and the isolation of habitats within mountains (so‐called “mountain islands”). However, parallels rarely go much beyond the observation that mountaintops are isolated from one another, as are true islands. Here, we challenge the analogy between mountains and true islands by re‐evaluating the literature, focusing on isolation (the prime mechanism underlying species endemism by restricting gene flow) from a dynamic perspective over space and time. Framework We base our conceptualization of “isolation” on the arguments that no biological system is completely isolated; instead, isolation has multiple spatial and temporal dimensions relating to biological and environmental processes. We distinguish four key dimensions of isolation: (a) environmental difference from surroundings; (b) geographical distance to equivalent environment [points (a) and (b) are combined as “snapshot isolation”]; (c) continuity of isolation in space and time; and (d) total time over which isolation has been present [points (c) and (d) are combined as “isolation history”]. We evaluate the importance of each dimension in different types of mountains and true islands, demonstrating that substantial differences exist in the nature of isolation between and within each type. In particular, different types differ in their initial isolation and in the dynamic trajectories they follow, with distinct phases of varying isolation that interact with species traits over time to form present‐day patterns of endemism. Conclusions Our spatio‐temporal definition of isolation suggests that the analogy between true islands and mountain islands masks important variation of isolation over long time‐scales. Our understanding of endemism in isolated systems can be greatly enriched if the dynamic spatio‐temporal dimensions of isolation enter models as explanatory variables and if these models account for the trajectories of the history of a system.