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Globally, ultramafic outcrops are renowned for hosting floras with high levels of endemism, including plants with specialised adaptations such as nickel or manganese hyperaccumulation. Soils derived from ultramafic regoliths are generally nutrient-deficient, have major cation imbalances, and have concomitant high concentrations of potentially phytotoxic trace elements, especially nickel. The South and Southeast Asian region has the largest surface occurrences of ultramafic regoliths in the world, but the geoecology of these outcrops is still poorly studied despite severe conservation threats. Due to the paucity of systematic plant collections in many areas and the lack of georeferenced herbarium records and databased information, it is not possible to determine the distribution of species, levels of endemism, and the species most threatened. However, site-specific studies provide insights to the ultramafic geoecology of several locations in South and Southeast Asia. The geoecology of tropical ultramafic regions differs substantially from those in temperate regions in that the vegetation at lower elevations is generally tall forest with relatively low levels of endemism. On ultramafic mountaintops, where the combined forces of edaphic and climatic factors intersect, obligate ultramafic species and hyperendemics often occur. Forest clearing, agricultural development, mining, and climate change-related stressors have contributed to rapid and unprecedented loss of ultramafic-associated habitats in the region. The geoecology of the large ultramafic outcrops of Indonesia’s Sulawesi, Obi and Halmahera, and many other smaller outcrops in South and Southeast Asia, remains largely unexplored, and should be prioritised for study and conservation.

Plants adapted to special soil types are ideal for investigating evolutionary processes, including maintenance of intraspecific variation, adaptation, reproductive isolation, ecotypic differentiation, and the tempo and mode of speciation. Common garden and reciprocal transplant approaches show that both local adaptation and phenotypic plasticity contribute to edaphic (soil-related) specialization. Edaphic specialists evolve rapidly and repeatedly in some lineages, offering opportunities to investigate parallel evolution, a process less commonly documented in plants than in animals. Adaptations to soil features are often under the control of major genes and they frequently have direct or indirect effects on genes that contribute to reproductive isolation. Both reduced competitiveness and greater susceptibility to herbivory have been documented among some edaphic specialists when grown in ‘normal’ soils, suggesting that a high physiological cost of tolerance may result in strong divergent selection across soil boundaries. Interactions with microbes, herbivores, and pollinators influence soil specialization either by directly enhancing tolerance to extremes in soil conditions or by reducing gene flow between divergent populations. Climate change may further restrict the distribution of edaphic specialists due to increased competition from other taxa or, expand their ranges, if preadaptations to drought or other abiotic stressors render them more competitive under a novel climate

Gypsum soils occur around the world, mainly in arid regions. These harsh environments promote unusual flora with high degrees of endemism. Mexico has extensive gypsum outcrops, but their flora has been poorly studied. However, the highest species richness and endemism are expected to be concentrated in Mexico’s northern dry regions. To promote the study of this flora and its conservation, we estimate how well sampled it is, quantify species richness, identify centers of endemism, and detect which gypsum outcrops lie within federal protected natural areas (PNA). We conducted exhaustive literature and herbaria reviews to generate a database of botanical records on gypsum soils. The total species and gypsophyte richness were calculated using cell grids. Centers of endemism were identified using the corrected weighted endemism index (CWE). We mapped the gypsum outcrops within PNA polygons. The most collected sites are Cuatro Ciénegas (Coahuila) and Santo Domingo Tonalá (Oaxaca), which also had the highest total species richness. Nevertheless, gypsophyte richness was higher in Cuatro Ciénegas and Nuevo León. The CWE identified seven gypsophyte centers of endemism. Mexico hosts the most diverse gypsophile flora in the world, despite having been only partially studied and collected. The regions with the highest species richness and endemism are unprotected.

Accurate census is essential for endangered plant management, yet lack of resources may make complete on-the-ground census difficult to achieve. Accessibility, especially for species in fragile habitats, is an added constraint. We examined the feasibility of using UAV (unmanned aerial vehicle, drone)-based imagery for census of an endangered plant species, Arctomecon humilis (dwarf bear-poppy), an herbaceous perennial gypsophile endemic of the Mojave Desert, USA. Using UAV technology, we captured imagery at both 50-m altitude (census) and 15-m altitude (validation) at two populations, White Dome (325 ha) and Red Bluffs (166 ha). The imagery was processed into orthomosaics that averaged 2.32 cm ground sampling distance (GSD) for 50-m imagery and 0.73 cm GSD for 15-m imagery. Putative poppy plants were marked in the 50-m imagery according to predefined criteria. We then used the 15-m imagery from each area to verify the identification accuracy of marked plants. Visual evaluation of the 50-m imagery resulted in errors of both commission and omission, mainly caused by failure to accurately identify or detect small poppies (<10 cm diameter). Higher-resolution 30-m altitude imagery (1.19 cm GSD) greatly reduced errors of commission. Habitat classification demonstrated that poppy density variation was closely tied to soil surface color. This study showed that drone imagery can potentially be used to census rare plant species with distinctive morphology in open habitats and understand their spatial distribution.

BACKGROUND: The Palaeotropical pitcher plant genus Nepenthes (Nepenthaceae) is characterized by specialized nutrient sequestration strategies, narrow endemism, and a patchy distribution in which vicariance is believed to have played a fundamental role. SCOPE: Using recent studies of the effects of climate, soil type and vicariance, we review patterns of diversity and endemism in Nepenthes. First we consider how climate influences the geographical range of the genus and diversity of prey trapping mechanisms. Second, we examine edaphic influences, specifically the relationship between limestone and ultramafic soils and the obligate edaphic endemic Nepenthes that inhabit them. Third, we examine the role of vicariance, with regards to the patchy distribution of suitable habitats throughout Southeast Asia, and the passive dispersal mechanism of Nepenthes seeds. CONCLUSIONS: Climate is the principal determinant of variation in pitcher functional traits and in perhumid environments, may drive the evolution of alternative nutrient sequestration strategies. Although little is known about the ecophysiological relationships between soil type and obligate edaphic Nepenthes, ultramafic and limestone substrates may strongly influence vegetation physiognomy, creating a diversity of environmental niches that are exploited by specialized Nepenthes species. Finally, the complex geology and geography of the Malay Archipelago drives diversification through vicariance.

The Cape Floristic Region (CFR) is globally recognized as a hotspot of plant diversity and endemism. Much of this diversity stems from radiations associated with infertile acid sands derived from sandstones of the geologically ancient Cape Fold Belt. These ancient montane floras acted as the source for most subsequent radiations on the Cape lowlands during the Oligocene (on silcretes) and Mio–Pliocene (on shales). The geomorphic evolution of the CFR during the Plio–Pleistocene led to the first large-scale occurrence of calcareous substrata (coastal dunes and calcarenites) along the Cape coast, providing novel habitats for plant colonization and ensuing evolution of the Cape coastal flora—the most recent diversification event in the Cape. Few studies have investigated the CFR’s dune and calcarenite floras, and fewer still have done so in an evolutionary context. Here, we present a unified flora of these coastal calcareous habitats of the CFR and analyze the taxonomic, biological and geographical traits of its component species to gain insights into its assembly. The Cape coastal flora, comprising 1,365 species, is taxonomically dominated by the Asteraceae, Fabaceae and Iridaceae, with Erica , Aspalathus and Agathosma being the most speciose genera. In terms of growth-form mix, there is a roughly equal split between herbaceous and woody species, the former dominated by geophytes and forbs, the latter by dwarf and low shrubs. Species associated with the Fynbos biome constitute the bulk of the flora, while the Subtropical Thicket and Wetland biomes also house a substantial number of species. The Cape coastal flora is a distinctly southern African assemblage, with 61% of species belonging to southern African lineages (including 35% of species with Cape affinity) and 59% being endemic to the CFR. Unique among floras from the Cape and coastal Mediterranean-climate regions is the relatively high proportion of species associated with tropical lineages, several of which are restricted to calcareous substrata of the CFR. The endemic, calcicolous component of the flora, constituting 40% of species, represents 6% of the Cape’s regional plant diversity—high tallies compared to other biodiversity hotspots. Most coastal-flora endemics emerged during the Plio–Pleistocene as a product of ecological speciation upon the colonization of calcareous substrata, with the calcifugous fynbos floras of montane acid substrata being the most significant source of this diversification, especially on the typically shallow soils of calcarenite landscapes. On the other hand, renosterveld floras, associated with edaphically benign soils that are widespread on the CFR lowlands, have not been a major source of lineages to the coastal flora. Our findings suggest that, over and above the strong pH gradient that exists on calcareous substrata, soil depth and texture may act as important edaphic filters to incorporating lineages from floras on juxtaposed substrata in the CFR.

QUESTIONS: Does edaphic heterogeneity (i.e. switches between gypsum and calcareous soils) affect annual plant community assembly in highly restrictive soils? Our main hypothesis is that soil filtering is the main determinant of species assembly, subsequently modulated by climate and lastly by biotic interactions, such as Stipa tenacissima and the biological soil crust (BSC) at fine spatial scales. Our study system is highly suitable to test the importance of edaphic heterogeneity to the assembly of annual plant communities, since calcareous and gypsum soils are in close contact and freely receive propagules. LOCATION: Annual plant communities of semi‐arid steppes in central Spain. METHODS: We built a soil affinity index (SAI) for each species to measure mean soil affinity (MSA; analogous to the community‐weighted mean – CWM) in each local assemblage. RESULTS: Most species were able to establish in both substrates, but gypsum soils clearly exerted a stronger filtering effect. Stipa favoured an expansion of the niche space in calcareous soils but not in gypsum, and BSCs not only reduced annual species richness, diversity and cover, but also the range of SAI values of local assemblages. CONCLUSIONS: This study highlights the importance of the probabilistic filtering (sensu From plant traits to vegetation structure, 2010, Cambridge University Press, Cambridge, UK) of soil characteristics to the assembly of annual plant communities, as opposed to an ‘all‐or‐nothing’ filter.

AIM: Granite outcrops may be able to act as refugia for species during adverse climate change, owing to their topographic complexity. We assessed this hypothesis by examining phylogeographical patterns in a common, geographically widespread granite endemic, Stypandra glauca (Hemerocallidaceae). LOCATION: Granite outcrops of the Southwest Australian Floristic Region, Western Australia. METHODS: Twenty‐four tetraploid individuals of the granite endemic Stypandra glauca were sampled from each of 12 granite outcrops: 7 from a mesic environment and 5 from the semi‐arid region. Phylogenetic reconstruction and divergence‐dating was achieved using Bayesian and parsimony analyses of chloroplast haplotypes from 90 individuals. Nuclear diversity and population differentiation were analysed across all individuals using 10 microsatellite loci. RESULTS: Stypandra glauca exhibited high (chloroplast) or moderate (nuclear) levels of divergence among, and low diversity within, outcrops. Haplotype diversity was high in both sampling regions, and each haplotype was unique to one outcrop. There was little correlation between geographical and genetic distance. Both nuclear and chloroplast diversity were higher in southern (mesic) outcrops than in northern (semi‐arid) outcrops, although the level of chloroplast divergence among outcrops was similar for both climatic regions. MAIN CONCLUSIONS: The levels of divergence and low diversity revealed in S. glauca support a scenario of prolonged isolation and persistence on granite outcrops in both mesic and semi‐arid climatic regions, with no evidence of contraction–expansion dynamics across the outcrop network. The higher levels of diversity in the southern populations may result from the maintenance of a larger effective population size in southern regions, which retained more mesic climates during drier glacial periods. Although the climatic conditions differ between outcrops in this study, our results indicate that outcrops in both regions have harboured S. glauca throughout climatic changes, accentuating the value of these habitats to biodiversity conservation under future changing climate.

BACKGROUND AND AIMS: Plants from gypsum habitats are classified as gypsophiles and gypsovags. The former include both narrow endemics limited to small gypsum areas and regionally dominant gypsophiles growing in gypsum areas of large regions, whereas gypsovags are plants that can grow both in gypsum and non-gypsum soils. Factors controlling the distribution of gypsum plants are still not fully understood. METHODS: To assess how the different types of gypsum plants deal with the stressful conditions of gypsum substrates, comparisons were made of the leaf chemical composition of four gypsovags, five regionally dominant gypsophiles and four narrow gypsum endemics growing in two massive gypsum areas of the Iberian Peninsula. KEY RESULTS: The chemical composition of gypsovags was clearly different from regionally dominant gypsophiles, while the chemical composition of narrow-gypsophile endemics was more similar to the chemical composition of gypsovags than to that of regionally dominant gypsophiles. Regionally dominant gypsophiles showed higher concentrations of ash, Ca, S, N, Mg P and Na, whereas gypsovags and local gypsophile endemics displayed higher concentrations of C and greater C : N ratios. CONCLUSIONS: Such differences suggest that the three groups of gypsum plants follow diverse ecological strategies. It is suggested that regionally dominant gypsophiles might fit the 'specialist' model, being species specifically adapted to gypsum, whereas both gypsovags and narrow-gypsophile endemics might fit the 'refuge' model, being stress-tolerant species that find refuge on gypsum soils from competition. The analysis of the leaf chemical composition could be a good predictor of the degree of plants specialization to gypsum soils.

Abstract Previous studies found that plant communities on infertile soils are relatively resistant to climatic variation due to stress tolerance adaptations. However, the species assemblies in gypsum soil habitats require further investigation. Thus, we considered the following questions. (1) Do harsher arid conditions determine the characteristics of the species that form plant assemblages? (2) Is the selection of the species that assemble in arid conditions mediated by their ability to grow on gypsum soils? (3) Is the selection of species that assemble in harsher conditions related to phylogenetically conserved functional traits? Perennial plant communities were analysed in 89 gypsum-soil sites along a 400 km climate gradient from the central to southeastern Iberian Peninsula. Each local assemblage was analysed in 30 × 30 m plots and described based on taxonomic, functional (soil plant affinity) and phylogenetic parameters. The mean maximum temperatures in the hottest month, mean annual precipitation and their interaction terms were used as surrogates for the aridity conditions in generalized linear models. In the hottest locations, the gypsophily range narrowed and the mean gypsophily increased at the community level, thereby suggesting the filtering of species and the dominance of soil specialists in the actual plant assemblies. Drier sites had higher taxonomic diversity. The species that formed the perennial communities were close in evolutionary terms at the two ends of the aridity gradient. The mean maximum temperatures in the hottest month had the main abiotic filtering effect on perennial plant communities, which was mediated by the ability of species to grow on gypsum soils, and thus gypsum specialists dominated the species assemblies in the hottest locations. In contrast, the perennial communities on gypsum soils were relatively resistant to changes in precipitation. Our findings suggest that the warmer environmental conditions predicted by global change models will favour gypsum specialists over generalists. We found that the perennial communities on gypsum soils were relatively resistant to changes in precipitation; however, temperatures in the hottest month were the main factor responsible for the selection of the species that finally established to form communities. It should be noted that species adapted to grow on gypsum soils (i.e. gypsophites) dominated plant communities in the hottest locations. Our findings suggest that the warmer environmental conditions predicted by global change models will favour gypsum specialists over generalists.