Rooting depth and xylem vulnerability are independent woody plant traits jointly selected by aridity, seasonality, and water table depth.

Evolutionary radiations of woody taxa within arid environments were made possible by multiple trait innovations including deep roots and embolism-resistant xylem, but little is known about how these traits have coevolved across the phylogeny of woody plants or how they jointly influence the distribution of species. We synthesized global trait and vegetation plot datasets to examine how rooting depth and xylem vulnerability across 188 woody plant species interact with aridity, precipitation seasonality, and water table depth to influence species occurrence probabilities across all biomes. Xylem resistance to embolism and rooting depth are independent woody plant traits that do not exhibit an interspecific trade-off. Resistant xylem and deep roots increase occurrence probabilities in arid, seasonal climates over deep water tables. Resistant xylem and shallow roots increase occurrence probabilities in arid, nonseasonal climates over deep water tables. Vulnerable xylem and deep roots increase occurrence probabilities in arid, nonseasonal climates over shallow water tables. Lastly, vulnerable xylem and shallow roots increase occurrence probabilities in humid climates. Each combination of trait values optimizes occurrence probabilities in unique environmental conditions. Responses of deeply rooted vegetation may be buffered if evaporative demand changes faster than water table depth under climate change.

The effects of foundation species on community assembly: a global study on alpine cushion plant communities.

Foundation species can change plant community structure by modulating important ecological processes such as community assembly, yet this topic is poorly understood. In alpine systems, cushion plants commonly act as foundation species by ameliorating local conditions. Here, we analyze diversity patterns of species' assembly within cushions and in adjacent surrounding open substrates (83 sites across five continents) calculating floristic dissimilarity between replicate plots, and using linear models to analyze relationships between microhabitats and species diversity. Floristic dissimilarity did not change across biogeographic regions, but was consistently lower in the cushions than in the open microhabitat. Cushion plants appear to enable recruitment of many relatively stress-intolerant species that otherwise would not establish in these communities, yet the niche space constructed by cushion plants supports a more homogeneous composition of species than the niche space beyond the cushion's influence. As a result, cushion plants support higher α-diversity and a larger species pool, but harbor assemblies with lower ß-diversity than open microhabitats. We conclude that habitats with and without dominant foundation species can strongly differ in the processes that drive species recruitment, and thus the relationship between local and regional species diversity.

Facilitative plant interactions and climate simultaneously drive alpine plant diversity.

Interactions among species determine local-scale diversity, but local interactions are thought to have minor effects at larger scales. However, quantitative comparisons of the importance of biotic interactions relative to other drivers are rarely made at larger scales. Using a data set spanning 78 sites and five continents, we assessed the relative importance of biotic interactions and climate in determining plant diversity in alpine ecosystems dominated by nurse-plant cushion species. Climate variables related with water balance showed the highest correlation with richness at the global scale. Strikingly, although the effect of cushion species on diversity was lower than that of climate, its contribution was still substantial. In particular, cushion species enhanced species richness more in systems with inherently impoverished local diversity. Nurse species appear to act as a 'safety net' sustaining diversity under harsh conditions, demonstrating that climate and species interactions should be integrated when predicting future biodiversity effects of climate change.

Underestimated diversity in high elevations of a global biodiversity hotspot: two new endemic species of Aethionema (Brassicaceae) from the alpine zone of Iran.

Although the mountains in South-West Asia are a global biodiversity hotspot, our understanding of their biodiversity, especially in the commonly remote alpine and subnival zones, is still limited. This is well exemplified here by Aethionema umbellatum (Brassicaceae), a species considered to have a wide yet disjoint distribution in the Zagros and Yazd-Kerman mountains of western and central Iran. Morphological and molecular phylogenetic data (based on plastid trnL-trnF and nuclear ITS sequences) show that A. umbellatum is restricted to a single mountain range in southwestern Iran (Dena Mts., southern Zagros), whereas populations from central Iran (Yazd-Kerman and central Zagros) and from western Iran (central Zagros) belong to species new to science, A. alpinum and A. zagricum, respectively. Both new species are phylogenetically and morphologically close to A. umbellatum, with which they share unilocular fruits and one-seeded locules. However, they are easily distinguishable by leaf shape, petal size, and fruit characters. This study confirms that the alpine flora of the Irano-Anatolian region is still poorly known. As the proportion of rare and local endemic species in alpine habitats is high, these habitats are of prime interest for conservation efforts.

Endemics determine bioregionalization in the alpine zone of the Irano-Anatolian biodiversity hotspot (South-West Asia).

Alpine habitats are characterized by a high rate of range restricted species compared to those of lower elevations. This is also the case for the Irano-Anatolian global biodiversity hotspot in South-West Asia, which is a mountainous area harbouring a high amount of endemic species. Using two quantitative approaches, Endemicity Analysis and Network-Clustering, we want to identify areas of concordant species distribution patterns in the alpine zone of this region as well as to test the hypothesis that, given the high proportion of endemics among alpine species, delimitation of these areas is determined mainly by endemic alpine species, i.e., areas of concordant species distribution patterns are congruent with areas of endemism. Endemicity Analysis identified six areas of concordant species distribution patterns irrespective of dataset (total alpine species versus endemic alpine species), whereas the Network-Clustering approach identified five and four Bioregions from total alpine species and endemic alpine species, respectively. Most of these areas have been previously identified using the endemic flora of different elevational zones. The identified units using both methods and both datasets are strongly congruent, proposing that they reveal meaningful distribution patterns. Bioregionalization in the Irano-Anatolian biodiversity hotspot appears to be strongly influenced by the endemic alpine species, a pattern likely to hold in alpine regions outside the Irano-Anatolian hotspot. SUPPLEMENTARY INFORMATION: The online version contains supplementary material available at 10.1007/s00035-021-00266-7.

The new locally endemic genus Yazdana (Caryophyllaceae) and patterns of endemism highlight the high conservation priority of the poorly studied Shirkuh Mountains (central Iran).

Although mountain ranges are often recognized as global biodiversity hotspots with a high level of endemism, diversity and biogeographic connections of isolated and weakly explored mountains remain poorly understood. This is also the case for Shirkuh Mts. in central Iran. Here, Yazdana shirkuhensis gen. & spec. nov. (Caryophylleae, Caryophyllaceae) is described and illustrated from the high alpine zone of this mountain. Molecular phylogenetic analyses of nuclear and plastid DNA sequence data show that Y. shirkuhensis is related to Cyathophylla and Heterochroa (tribe Caryophylleae). The newly described genus and species accentuate Shirkuh Mts. as a center of endemism, which harbors a high number of narrowly distributed species, mostly in high elevations reaching alpine habitats. As this area is currently not protected, a conservation priority is highlighted for high elevations of Shirkuh Mts.

Endemic diversity and distribution of the Iranian vascular flora across phytogeographical regions, biodiversity hotspots and areas of endemism.

Endemism is one of the most important concepts in biogeography and is of high relevance for conservation biology. Nevertheless, our understanding of patterns of endemism is still limited in many regions of high biodiversity. This is also the case for Iran, which is rich in biodiversity and endemism, but there is no up-to-date account of diversity and distribution of its endemic species. In this study, a comprehensive list of all endemic vascular plant species of Iran, their taxonomic composition and their geographical distribution are presented. To this end, a total of 2,597 (sub)endemic vascular plant species of Iran were documented and their distribution in three phytogeographical regions, two biodiversity hotspots and five areas of endemism were analysed. The Irano-Turanian phytogeographical region harbours 88% of the Iranian endemics, the majority of which are restricted to the Irano-Anatolian biodiversity hotspot (84%). Nearly three quarters of the endemic species are restricted to mountain ranges. The rate of endemism increases along an elevational gradient, causing the alpine zone to harbour a disproportionally high number of endemics. With increasing pastoralism, urbanization, road construction and ongoing climate change, the risk of biodiversity loss in the Iranian mountains is very high, and these habitats need to be more effectively protected.

A bioclimatic characterization of high elevation habitats in the Alborz mountains of Iran.

The Alborz mountains in N-Iran at 36° N rise from the Caspian Sea to 5671 m a.s.l., with warm-temperate, winter-deciduous forests in the lower montane belt in northern slopes, and vast treeless terrain at higher elevation. A lack of rainfall (ca. 550 mm at high elevations) cannot explain the absence of trees. Hence, it is an open question, which parts of these mountains belong to the alpine belt. Here we use bioclimatic data to estimate the position of the potential climatic treeline, and thus, define bioclimatologically, what is alpine and what is not. We employed the same miniature data loggers and protocol that had been applied in a Europe-wide assessment of alpine climates and a global survey of treeline temperatures. The data suggest a potential treeline position at ca. 3300 m a.s.l., that is ca. 900 m above the upper edge of the current oak forest, or 450 m above its highest outposts. The alpine terrain above the climatic treeline position shows a temperature regime comparable to sites in the European Alps. At the upper limit of angiosperm life, at 4850 m a.s.l., the growing season lasted 63 days with a seasonal mean root zone temperature of 4.5 °C. We conclude that (1) the absence of trees below 2850 m a.s.l. is clearly due to millennia of land use. The absence of trees between 2850 and 3300 m a.s.l. is either due to the absence of suitable tree taxa, or the only potential regional taxon for those elevations, Juniperus excelsa, had been eradicated by land use as well. (2) These continental mountains provide thermal life conditions in the alpine belt similar to other temperate mountains. (3) Topography and snow melt regimes play a significant role for the structure of the alpine vegetation mosaics.

Hotspots within a global biodiversity hotspot - areas of endemism are associated with high mountain ranges.

Conservation biology aims at identifying areas of rich biodiversity. Currently recognized global biodiversity hotspots are spatially too coarse for conservation management and identification of hotspots at a finer scale is needed. This might be achieved by identification of areas of endemism. Here, we identify areas of endemism in Iran, a major component of the Irano-Anatolian biodiversity hotspot, and address their ecological correlates. Using the extremely diverse sunflower family (Asteraceae) as our model system, five consensus areas of endemism were identified using the approach of endemicity analysis. Both endemic richness and degree of endemicity were positively related to topographic complexity and elevational range. The proportion of endemic taxa at a certain elevation (percent endemism) was not congruent with the proportion of total surface area at this elevation, but was higher in mountain ranges. While the distribution of endemic richness (i.e., number of endemic taxa) along an elevational gradient was hump-shaped peaking at mid-elevations, the percentage of endemism gradually increased with elevation. Patterns of endemic richness as well as areas of endemism identify mountain ranges as main centres of endemism, which is likely due to high environmental heterogeneity and strong geographic isolation among and within mountain ranges. The herein identified areas can form the basis for defining areas with conservation priority in this global biodiversity hotspot.