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Background and Aims The leaf economic spectrum (LES) is related to dry mass and nutrient investments towards photosynthetic processes and leaf structures, and to the duration of returns on those investments (leaf lifespan, LL). Phosphorus (P) is a key limiting nutrient for plant growth, yet it is unclear how the allocation of leaf P among different functions is coordinated with the LES. We addressed this question among 12 evergreen woody species co-occurring on P-impoverished soils in south-eastern Australia. Methods Leaf ‘economic’ traits, including LL, leaf mass per area (LMA), light-saturated net photosynthetic rate per mass (Amass), dark respiration rate, P concentration ([Ptotal]), nitrogen concentration, and P resorption, were measured for three pioneer and nine non-pioneer species. Leaf P was separated into five functional fractions: orthophosphate P (Pi), metabolite P (PM), nucleic acid P (PN), lipid P (PL), and residual P (PR; phosphorylated proteins and unidentified compounds that contain P). Results LL was negatively correlated with Amass and positively correlated with LMA, representing the LES. Pioneers occurred towards the short-LL end of the spectrum and exhibited higher [Ptotal] than non-pioneer species, primarily associated with higher concentrations of Pi, PN and PL. There were no significant correlations between leaf P fractions and LL or LMA, while Amass was positively correlated with the concentration of PR. Conclusions Allocation of leaf P to different fractions varied substantially among species. This variation was partially associated with the LES, which may provide a mechanism underlying co-occurrence of species with different ecological strategies under P limitation.

To date, there have been few studies of the functional traits of the dioecious Hippophae tibetana Schlecht leaves, either male or female, in response to ecological factors such as altitude. Elucidating these relationships will establish an important scientific basis for vegetation restoration and reconstruction of the Tibetan Plateau ecosystem. The natural populations of H. tibetana, distributed across three field sites, at 2868 m, 3012 m and 3244 m, in Tianzhu, Gansu, were studied by field survey sampling and laboratory analysis. In particular, the adaptions of leaf functional traits to elevation in these dioecious plants were analyzed. The results show that: (1) there is no “midday depression” of photosynthetic activity in either male or female plants. Over a one-day period, the net photosynthetic rate (Pn) and transpiration rate (Tr) of H. tibetana female plants were higher than those of male plants (p < 0.05). This correlated to the period of vigorous fruit growth in the female plant. The measured Pn and Tr were maximal at the intermediate altitude (3012 m). The light compensation point (LCP) of the leaves of male and female plants were 57.6 and 43.2 μmol·m−2·s−1, respectively, and the light saturation points (LSP) of the leaves were 1857.6 and 1596.8 μmol·m−2·s−1. (2) Altitude had a significant effect on plant and leaf functional traits of male and female H. tibetana (p < 0.05), and no significant difference was noted between plants at the same altitude. The values for leaf area (LA), specific leaf weight (LMA), leaf phosphorus content per unit mass (Pmass) and leaf phosphorus content per unit area (Parea) were also maximal at the intermediate altitude. Leaf nitrogen content per unit area (Narea) and leaf nitrogen content per unit mass (Nmass) increased with altitude. This indicated that the functional traits of male and female plants and leaves of H. tibetana showed a strong “trade-off relationship” with altitude. (3) Pearson correlation analysis showed that there were significant correlations among functional traits of H. tibetana leaves. Redundancy analysis (RDA) showed that soil water content (SWC), altitude (Alt) and soil organic carbon (SOC) had significant effects on the functional traits of H. tibetana leaves (p < 0.05).

Calligonum mongolicum is a dominant native perennial shrub on active sand dunes in arid deserts of Northern China, and is therefore widely used in sand dune stabilisation in these regions. However, it remains largely unknown about how seedling emergence of C. mongolicum has adapted to unpredictable sand movement and extreme drought. Here we examined effects of sand burial depth, light intensity and seed age on seedling emergence, and consider seed germination and seedling emergence strategies for the shrubs adaption to the desert environment. In our pot experiment, the optimum seeding depth for emergence of C. mongolicum was 2 cm, indicating that for germination and seedling emergence only moderate sand burial is required. Light intensity at the surface soil (0 cm) was important for seedling emergence, while there was no significant difference between 50 and 20 % light flux density, at burial depths of 1 and 2 cm, indicating that C. mongolicum seeds had adapted to sand burial, while not sand erosion. We also found C. mongolicum seedlings emerged in Spring, and also in late Summer to early Autumn. Meanwhile, the final percentage of seedling emergence for 3-year-old seeds, was similar to that of 1-year-old seeds, which meant that C. mongolicum seeds were well conserved under natural conditions in sand dunes, thus were capable of developing a persistent but shallow soil seed-bank. The results indicate that germination and seedling emergence take a bet-hedging strategy to adapt to the extreme desert environment. Our study clarified that the C. mongolicum desert shrub has combined strategies in its adaption to arid sand environments.

Sand burial and wind erosion caused by sand movement are common phenomena in desert environments, but the effects on clonal shrub have rarely been investigated. Here, we assessed how sand movements affect the population regeneration capacity of juvenile clonal fragments of the shrub Calligonum mongolicum growing in mobile desert sand dunes. We investigated the population status and natural regeneration capacity in three types of mobile dunes (heavy wind erosion, heavy sand burial and moderate sand burial). Clonal propagation of C. mongolicum was markedly different across sites. Moderate sand burial sites had the largest ramet density and bud number per unit length of rhizome, and the overwinter survival rate was significantly higher at sand burial sites than at wind erosion sites, suggesting that C. mongolicum may have well adapted to the moderate sand burial environment. We further examined the effects of clonal integration on clonal regeneration of this species. Physiological, biochemical and morphological characteristics of parent and daughter ramets growing in heterogeneous sandy habitats (sand burial or wind erosion) were measured. The results showed that being connected or severed from the maternal plant critically determined survival of daughter ramets on wind eroded rhizomes. When eroded rhizomes remained connected, the mother ramets had the highest chlorophyll a, b and a + b contents. However, both the mother plant and the daughter ramets undergoing erosion had higher proline and soluble protein levels than sand buried ramets. Meanwhile, the daughter ramets undergoing sand burial had higher photosynthetic rates (Pn), chlorophyll fluorescence parameters (Fm and Fo), and phenotypic traits of assimilating shoots, i.e., node number, length and volume than wind-eroded ramets. However, significant differences with mother plants, whether connected or severed, were very limited. It was concluded that moderate sand burial environments promoted clonal reproduction and growth of C. mongolicum. Additionally, physiological integration with mother raments in favorable conditions can alleviate stress on daughter ramets exposed to wind erosion. This physiological effect may do not occur for sand buried daughter ramets. These survival strategies and phenotypic responses should be carefully considered in shrub and sand dune management in sand fixation plantations of C. mongolicum.

Calligonum mongolicum is a dominant native perennial shrub on sand dunes in arid deserts of northwestern China, and is therefore widely used in sand dune stabilization in these regions. However, it remains largely unknown how seedling emergence of C. mongolicum has adapted to unpredictable sand movement and extreme drought. Here we examined effects of seed burial depth, light intensity, and seed age on seedling emergence, and considered seed germination and seedling emergence strategies for the shrub’s adaption to the desert environment. In our pot experiment, the optimum seeding depth for emergence of C. mongolicum was 2 cm, indicating that for germination and seedling emergence only moderate sand burial is required. Light intensity at the surface soil (0 cm) was important for seedling emergence, while there was no significant difference between 50 and 20% light flux density, at burial depths of 1 and 2 cm, indicating that C. mongolicum seeds had adapted to sand burial, while not exposure from sand erosion. We also found C. mongolicum seedlings emerged in spring and in late summer to early autumn. Meanwhile, seedling emergence percentage for 3-year-old seeds was similar to that of 1-year-old seeds, which meant that C. mongolicum seeds were well preserved under normal sand dune conditions, thus were capable of developing a persistent, but shallow soil seed-bank. These results indicated that germination and seedling emergence take a bet-hedging strategies to adapt to variable desert environments. Our study confirmed that C. mongolicum desert shrubs combine strategies in its adaption to arid and variable sand environments.