Serpentine soils limit plant growth by NPK deficiencies, low Ca availability, excess Mg, and high heavy metal levels. In this study, three congeneric serpentine and nonserpentine evergreen shrub species pairs were grown in metalliferous serpentine soil with or without NPKCa fertilizer to test which soil factors most limit biomass production and mineral nutrition responses. Fertilization increased biomass production and allocation to leaves while decreasing allocation to roots in both serpentine and nonserpentine species. Simultaneous increases in biomass and leaf N:P ratios in fertilized plants of all six species suggest that N is more limiting than P in this serpentine soil. Neither N nor P concentrations, however, nor root to shoot translocation of these nutrients, differed significantly between serpentine and nonserpentine congeners. All six species growing in unfertilized serpentine soil translocated proportionately more P to leaves compared to fertilized plants, thus maintaining foliar P. Leaf Ca:Mg molar ratios of the nonserpentine species were generally equal to that of the soil. The serpentine species, however, maintained significantly higher leaf Ca:Mg than both their nonserpentine counterparts and the soil. Elevated leaf Ca:Mg in the serpentine species was achieved by selective Ca transport and/or Mg exclusion operating at the root-to-shoot translocation level, as root Ca and Mg concentrations did not differ between serpentine and nonserpentine congeners. All six species avoided shoot toxicity of heavy metals by root sequestration. The comparative data on nutrient deficiencies, leaf Ca:Mg, and heavy metal sequestration suggest that the ability to maintain high leaf Ca:Mg is a key evolutionary change needed for survival on serpentine soil and represents the physiological feature distinguishing the serpentine shrub species from their nonserpentine congeners. The results also suggest that high leaf Ca:Mg is achieved in these serpentine species by selective translocation of Ca and/or inhibited transport of Mg from roots, rather than by uptake/exclusion at root surfaces.
Although several lichen inventories exist for European ultramafic sites, only four surveys of serpentine lichens for North America have been published to date. Of those, only one has been conducted in California. We conducted a survey of saxicolous lichens from ultramafic rocks (including nephrite, partially serpentinized peridotite, and serpentinite) and non-ultramafic rocks (including silica-carbonate, shale, and sandstone) at the New Idria serpentinite mass, San Benito County, California. X-ray Fluorescence Analysis of the rocks from which the lichens were collected revealed significant elemental differences between the ultramafic and non-ultramafic rocks for 26 of the 32 major and trace elements analyzed. We identified a total of 119 species of lichenized and lichenicolous fungi; 60 species were restricted to ultramafic substrata, 19 to silica-carbonate, and 15 to shale and sandstone. Only 4 species were shared in common. A permutational multivariate analysis of variance (perMANOVA) test revealed significant differences in lichen assemblages between ultramafic and non-ultramafic rocks at the species level but not at the generic level, with species richness (alpha-diversity) significantly greater at the ultramafic sites. We suggest that, although differences in geochemistry clearly influence the lichen community composition, other factors, especially substratum age and the physical characteristics of the rock, are of equal, if not greater, importance. Of all the species collected, six, Buellia aethalea, B. ocellata, Caloplaca oblongula, Rhizocarpon saurinum, Thelocarpon laureri, and Trapelia obtegens, are reported new to California, along with an apparently previously undescribed Solenopsora sp. The rest of the species encountered are relatively frequent in the lichen flora of southern and central California, except Aspicilia praecrenata, a rare California endemic that we collected on both ultramafic and non-ultramafic rocks.
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