Invasive alien plants in marine protected areas: the <Emphasis Type="Italic">Spartina anglica</Emphasis> affair in the European Wadden Sea

The common cord-grass Spartina anglica, a fertile hybrid of S. maritima and S. alterniflora, was planted in the European Wadden Sea extensively during the late 1920s and 1930s to promote sediment accretion. After establishment, it colonised as a pioneer plant in the upper tidal zone, where it occurs frequently in coherent swards at the seaward front of saltmarshes and in patches on the tidal flats. Often, a conspicuous, almost monotypic, belt of S. anglica is formed. Over the last two decades, an increase in abundance and accelerated spread of S. anglica was observed, possibly promoted by warmer spring temperatures. This alien species may benefit from global warming, and there is considerable concern about its harmful impacts on the native biocoenoses and native biodiversity of the unique Wadden Sea ecosystem, encompassing effects on hydromorphodynamics and coastal protection. For a definitive assessment, however, an adequate quantification and comparison of documented and potential effects of S. anglica is important, but currently unavailable. Consequently, no management strategy exists for the prevention or restoration of the Wadden Sea ecosystem. Thus, the development of an alien species plan on the level of the Trilateral Cooperation on the Protection of the Wadden Sea is essential.     

Density-dependent effects on the dieback of exotic species Spartina anglica in coastal China

The exotic species Spartina anglica C.E. Hubbard has been experiencing dieback in coastal China during the past decade, while it is invading other places in the world. Thus, an explanation for the dieback of the populations in coastal China would be helpful for controlling invasions of S. anglica in other places of the world. Both field and greenhouse experiments were conducted to determine the possible role of density in the dieback of the populations in China. The density effects were tested by maintaining six density levels in field experiments and three density levels in greenhouse experiments over 7 months. The results showed that growth, asexual reproduction and biomass accumulation in medium density were significantly higher than those in other density treatments for field experiments. The growth and asexual reproduction in high-density patches were lowest among the treatments in the field. Data showed the same pattern for the greenhouse experiments. The results implied that density regulation had a great effect on most of the studied parameters. Because the species is surviving at high density along the Chinese coast, and has a much lower growth rate due to strong intraspecific competition, and also because high-density populations may decline into low density populations due to resource exhaustion, high plant density might be one of the important factors in the dieback of S. anglica.     

Shade and salinity responses of two dominant coastal wetland grasses: implications for light competition at the transition zone

BackgroundCoastal wetlands are threatened by the increased salinity that may result from sea level rise. Salinity stress alters species zonation patterns through changes in competitive outcome between species differing in salinity tolerance. This study therefore aimed to understand how salinity and light affect two dominant and competing coastal wetland grasses that differ in salt tolerance, height and photosynthetic metabolism.MethodsThe C₄ species Spartina anglica and the C₃ species Phragmites australis were grown at five salinity levels (0, 7, 14, 21 and 28 ppt) and two light fluxes (100 % and 50 % of natural daylight) in an outdoor experimental setup for 102 d with full access to nutrients.Key ResultsSalinity reduced the biomass, height and shoot density of P. australis from 81.7 g dry weight (DW), 0.73 m and 37 shoots per pot at a salinity of 0 ppt to 16.8 gDW, 0.3 m and 14 shoots per pot at a salinity of 28 ppt. Biomass, height and shoot density of S. anglica did not respond or were only slightly reduced at the highest salinity of 28 ppt. High salinity also resulted in a higher tissue concentration of N and P in P. australis. Both species had low ability to acclimate to the lower light flux. Shade acclimation in S. anglica occurred via modest changes in specific leaf area, pigment content and biomass allocation.ConclusionsHigh salinity reduced traits important for light competition and increased the nutrient concentration in P. australis leaf and root biomass, while this was overall unaffected in S. anglica. This is likely to reduce the competitive ability of P. australis over S. anglica for light because at high salinities the former cannot effectively shade the lower-growing S. anglica. Neither species effectively acclimates to shade, which could explain why S. anglica does not occur in the understorey of P. australis at low salinities.     

Remedial habitat creation: does Nereis diversicolor play a confounding role in the colonisation and establishment of the pioneering saltmarsh plant, Spartina anglica?

Increasing concerns over global warming and expected sea level rises have led to the adoption of new coastal management strategies around the south-east coast of England. This paper explores the role played by the estuarine invertebrate Nereis diversicolor in limiting the colonisation and establishment of the invasive pioneering salt marsh plant, Spartina anglica. The biology of N. diversicolor is briefly reviewed and data from field experiments are presented demonstrating significant negative effects of worm abundance on transplanted S. anglica biomass. Laboratory-based experiments demonstrated significant negative effects of N. diversicolor abundance on the survival of S. anglica seeds transplanted to sediment cores. The importance of estuarine invertebrates in engineering the mudflat habitat may confound the foreseen ecosystem services and function provided by saltmarsh management schemes. Received: 15 February 1999 / Received in revised form: 4 June 1999 / Accepted: 11 June 1999     

Limited seed retention during winter inhibits vegetation establishment in spring,affecting lateral marsh expansion capacity

Coastal systems worldwide deliver vital ecosystem services, such as biodiversity, carbon sequestration, and coastal protection. Effectivity of these ecosystem services increases when vegetation is present. Understanding the mechanisms behind vegetation establishment in bio‐geomorphic systems is necessary to understand their ability to recover after erosive events and potential adaptations to climate change. In this study, we examined how seed availability affects vegetation establishment in the salt marsh–intertidal flat transition zone: the area with capacity for lateral marsh expansion. This requires vegetation establishment; therefore, seed availability is essential. In a 6‐month field experiment, we simulated a before and after winter seed dispersal at two locations, the salt‐marsh vegetation edge and the intertidal flat, and studied seed retention, the seed bank, and the seed viability of three pioneer marsh species: Salicornia procumbens, Aster tripolium, and Spartina anglica. During winter storm conditions, all supplied seeds eroded away with the sediment surface layer. After winter, supplied seeds from all three species were retained, mostly at the surface while 9% was bioturbated downwards. In the natural seed bank, A. tripolium and S. anglica were practically absent while S. procumbens occurred more frequently. The viability of S. procumbens seeds was highest at the surface, between 80% and 90%. The viability quickly decreased with depth, although viable S. procumbens seeds occurred up to 15 cm depth. Only when seeds were supplied after winter, many S. procumbens and some S. anglica individuals did establish successfully in the transition zone. Viable seed availability formed a vegetation establishment threshold, even with a local seed source. Our results suggest that, although boundary conditions such as elevation, inundation, and weather conditions were appropriate for vegetation establishment in spring, the soil surface in winter can be so dynamic that it limits lateral marsh expansion. These insights can be used for designing effective nature‐based coastal protection.     

Eco-physiological responses of the declining population Spartina anglica to N and P fertilizer addition

Nitrogen and phosphorus are both important life elements. N, P and combined N-P fertilizers were added to the declining population Spartina anglica Hubbard in coastal China. Some growth parameters and eco-physiological responses of S. anglica to different fertilizer treatments (N, P and combined N-P fertilizer addition with high, medium and low levels, respectively) were measured. The fertilizer addition had a highly significant effect on the dynamics of its height-growth, number of leaves, number of roots and total biomass. Only N addition had a significant effect on leaf area and leaf thickness in all fertilizer treatments. On the dynamics of its height-growth, the effect of N addition was the most apparent, and the effect of N-P addition was not greater than those of N and P addition separately. The photosynthesis rate was enhanced and the yield was the highest with the highest N, the highest N-P and the medium P addition. The rates were higher than those of CK by 19.08 μmol·m^?2·s^?1, 15.47 μmol·m^?2·s^?1 and 11.23 μmol·m^?2·s^?1, respectively. The activity of SOD and POD increased with the treatments after freshwater stress for 14 days. Effects of medium N and P addition were significant for SOD activity. However, POD activity was significantly higher with the treatment of higher N and higher N-P addition. In a word, fertilizer addition improved the growth of the declining population S. anglica. The results indicated that the decline of S. anglica was correlated with the nutriment deficiency in soil, especially with the lack of N.     

Effects of sea level rise on mangrove Avicennia population growth,colonization and establishment: Evidence from a field survey and greenhouse manipulation experiment

Mangroves will either face longer tidal inundation or retreat landwards in response to on-going accelerated sea level rise. However, little is known about the growth, regeneration or colonization of mangrove seedlings under the different tidal inundation regimes associated with accelerated sea level rise. In the present study, a field survey and a greenhouse mesocosm experiment were conducted to evaluate possible effects of accelerated sea level rise on colonization, establishment and seedling growth of a mangrove pioneer species, Avicennia marina. Avicennia populations at different elevations of the intertidal zone on Xiamen Island in Fujian Province, China showed distinctive patterns in both seedling density and plant age. The seedlings at lower elevations had less annual biomass accumulation and population productivity, but higher shoot to root ratios, suggesting that elevation has positive effects on seedling growth. The greenhouse mesocosm experiment with 1-year-old A. marina seedlings utilized five inundation periods (0, 2, 4, 6 and 12 h in a semidiurnal tidal cycle) and two inundation depths (root immersion and canopy immersion). Both inundation period and depth exerted significant and negative effects on biomass accumulation, photosynthetic rate, leaf electron transportation and water use efficiency. However, the negative effects of canopy immersion were more profound than root immersion. Canopy immersion exacerbated the effects of prolonged inundation, with no seedlings surviving under canopy immersion at the 12-hour treatment. These results suggest that at lower elevations with higher sea level, canopy immersion will have greater negative effects on seedling colonization, establishment and early growth of A. marina. This finding is instrumental in predicting the future dynamics of mangrove forests under increasing sea levels.     

Morphological and biomechanical responses of floodplain willows to tidal flooding and salinity

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Photosynthetic and physiological responses of Kandelia candel L. Druce seedlings to duration of tidal immersion in artificial seawater

The present study demonstrates the influence of the duration of periodical waterlogging with artificial seawater on the photosynthetic and physiological responses of Kandelia candel L. Druce seedlings, the pre-dominant species of subtropical mangroves in China. Artificial tidal fluctuations applied here closely mimicked the twice daily tidal inundation which mangroves experience in the field. All the seedlings were immersed in artificial seawater during 70-day cultivation. Similar trends with increasing duration of immersion occurred in photosynthetic rate, transpiration rate, stomatal conductance and intercellular CO₂ concentration, where significant decreases occurred only in long time treatments of 10 or 12 h. Water used efficiency and chlorophyll contents showed lower in medium periods and higher in long periods of immersion. This indicates that the increase in pigment contents of leaves was ineffective in promoting P_n under long time immersion. Light saturation points under short time waterlogging (0–4 h) occurred at light intensities of 800–1000 μmol/m²/s, and at around 400 μmol/m²/s in long time treatments (8–12 h). Long periods of tidal immersion therefore significantly inhibited photosynthesis of mature leaves. Alcohol dehydrogenase and oxidase activity in roots both increased under longer immersion periods, suggesting that roots are sensitive to anaerobiosis under long term waterlogging. The activities of peroxidase and superoxide dismutase in mature leaves increased in 8 h and 10 h treatments, respectively. The content of malondialdehyde in mature leaves increased under long time treatments. Abscisic acid accumulation in mature leaves also had a sharp increase from 8 h to 12 h inundation. Even though the anti-oxidative enzymes were induced by waterlogging, this was not sufficient to protect the seedlings from senescence. The results suggested that K. candel seedlings completely tolerated tidal immersion by seawater up to about 8 h in each cycle, which matches the natural distribution of K. candel in inter-tidal zones of China.     

The sediment burial depth and salinity control the early developments of Suaeda salsa in the Yellow River Delta

Sediment deposition is a common phenomenon in the estuary area. Pot control experiments were conducted to evaluate the interaction effects of sediment burial depth and salt stress on the seed germination and early seedling growth of Suaeda salsa (L.) Pall., an pioneer species of tidal wetland near the Yellow River Delta. The results showed that the percentage of seedling emergence, seedling emergence rate, seedling height, branch number, shoot biomass and root biomass were all significantly affected by salt stress and sediment burial depth. While the interaction of salt and burial depth significantly influenced the branch number, leaf biomass, shoot biomass and total plant biomass. Only 5 cm burial depth without salt stress should 6.25 ± 3.61% seedlings emergence. With the increasing of sediment burial depth and salt stress, percentage of seedling emergence, seedling emergence rate and plant height decreased significantly. However, under the salt treatment of 0 and 1%, the branch number increased dramatically with the increasing of sediment burial depth from 0 to 3 cm. The ratio of leaf to total biomass increased with increasing of burial depth, on the contrary, the ratio of root to total biomass decreased. 0–1 cm sediment burial depth was proved the suitable depths for seed germination of S. salsa in the coastal wetland of the Yellow River Delta. Our findings contribute to a better understanding of how to improve the seedling establishment of S. salsa under the dynamic changes of sediment deposition and salinity in the coastal wetland of the Yellow River Delta.     

Cultivation of the brown alga Sargassum horneri: sexual reproduction and seedling production in tank culture under reduced solar irradiance in ambient temperature

As a large conspicuous intertidal brown alga, individuals of Sargassum horneri can reach a length of more than 7 m with a fresh weight of 3 kg along the coasts of the Eastern China Sea. The biomass of this alga as a vital component in coastal water ecology has been well documented. In recent years, a steady disappearance of the algal biomass along the once densely populated coastal areas of the Eastern China Sea has drawn attention in China. Efforts have been made to reconstruct the subtidal algal flora or even to grow the alga by use of long-lines. As part of the efforts to establish an efficient technique for producing seedlings of S. horneri, in this investigation a series of culture experiments were carried out in indoor raceway and rectangular tanks under reduced solar irradiance at ambient temperature in 2007–2008. The investigation demonstrated that: (1) sexual reproduction of S. horneri could be accelerated in elevated temperature and light climates, at least 3 months earlier than in the wild; (2) eggs of S. horneri had the potential to be fertilized up to 48 h, much longer than that of known related species; (3) suspension and fixed culture methods were both effective in growing the seedlings to the long-line cultivation stage; and (4) the life cycle of S. horneri in culture could be shortened to 4.5 months, thus establishing this alga as an appropriate model for investigating sexual reproduction in dieocious species of this genus.     

Phenotypic plasticity of four Chenopodiaceae species with contrasting saline–sodic tolerance in response to increased salinity–sodicity

It is unknown whether phenotypic plasticity in fitness‐related traits is associated with salinity–sodicity tolerance. This study compared growth and allocation phenotypic plasticity in two species with low salinity–sodicity tolerance (Chenopodium acuminatum and C. stenophyllum) and two species with high salinity–sodicity tolerance (Suaeda glauca and S. salsa) in a pot experiment in the Songnen grassland, China. While the species with low tolerance had higher growth and allocation plasticity than the highly tolerant species, the highly tolerant species only adjusted their growth traits and maintained higher fitness (e.g., plant height and total biomass) in response to increased soil salinity–sodicity, with low biomass allocation plasticity. Most plasticity is “apparent” plasticity (ontogenetic change), and only a few traits, for example, plant height:stem diameter ratio and root:shoot biomass ratio, represent “real” plasticity (real change in response to the environment). Our results show that phenotypic plasticity was negatively correlated with saline–sodic tolerance and could be used as an index of species sensitivity to soil salinity–sodicity.     

Nitrogen level changes the interactions between a native (Scirpus triqueter) and an exotic species (Spartina anglica) in Coastal China

The exotic species Spartina anglica, introduced from Europe in 1963, has been experiencing a decline in the past decade in coastal China, but the reasons for the decline are still not clear. It is hypothesized that competition with the native species Scirpus triqueter may have played an important role in the decline due to niche overlap in the field. We measured biomass, leaf number and area, asexual reproduction and relative neighborhood effect (RNE) of the two species in both monoculture and mixture under three nitrogen levels (control, low and high). S. anglica showed significantly lower biomass accumulation, leaf number and asexual reproduction in mixture than in monoculture. The inter- and intra-specific RNE of S. anglica were all positive, and the inter-specific RNE was significantly higher than the intra-specific RNE in the control. For S. triqueter, inter- and intra-specific RNE were negative at the high nitrogen level but positive in the control and at the low nitrogen level. This indicates that S. triqueter exerted an asymmetric competitive advantage over S. anglica in the control and low nitrogen conditions; however, S. anglica facilitated growth of S. triqueter in high nitrogen conditions. Nitrogen level changed the interactions between the two species because S. triqueter better tolerated low nitrogen. Since S. anglica is increasingly confined to upper, more nitrogen-limited marsh areas in coastal China, increased competition from S. triqueter may help explain its decline.     

Phenotypic plasticity of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> and <Emphasis Type="Italic">Phragmites australis</Emphasis> in response to nitrogen addition and intraspecific competition

Phenotypic plasticity of the two salt marsh grasses Spartina alterniflora and Phragmites australis in salt marshes is crucial to their invasive ability, but the importance of phenotypic plasticity, nitrogen levels, and intraspecific competition to the success of the two species is unclear at present. Spartina alterniflora Loisel. is an extensively invasive species that has increased dramatically in distribution and abundance on the Chinese and European coasts, and has had considerable ecological impacts in the regions where it has established. Meanwhile, Phragmites australis Cav., a native salt marsh species on the east coast of China, has replaced the native S. alterniflora in many marshes along the Atlantic Coast of the US. This study determined the effects of nitrogen availability and culm density on the morphology, growth, and biomass allocation traits of Spartina alterniflora and Phragmites australis. A large number of morphological, growth, and biomass parameters were measured, and various derived values (culm: root ratio, specific leaf area, etc.) were calculated, along with an index of phenotypic plasticity. Nitrogen addition significantly affected growth performance and biomass allocation traits of Spartina alterniflora, and culm density significantly affected morphological characteristics in a negative way, especially for Spartina alterniflora. However, there were no significant interactions between nitrogen levels and culm density on the morphological parameters, growth performances parameters, and biomass allocation parameters of the two species. Spartina alterniflora appears to respond more strongly to nitrogen than to culm density and this pattern of phenotypic plasticity appears to offer an expedition for successful invasion and displacement of Phramites australias in China. The implication of this study is that, in response to the environmental changes that are increasing nitrogen levels, the range of Spartina alterniflora is expected to continue to expand on the east coast of China.     

Different combinations of morpho‐physiological traits are responsible for tolerance to drought in wild tomatoes Solanum chilense and Solanum peruvianum

Herbaceous species can modify leaf structure during the growing season in response to drought stress and water loss. Evolution can select combinations of traits in plants for efficient water use in restricted environments. We investigated plant traits that mediate adaptation and acclimation to water stress in two herbaceous drought‐tolerant species. Anatomical, morphological and physiological traits related to stems and leaves were examined under optimal watering (OW) and a long period of restricted watering (RW) in 11 accessions from three Solanaceae species (Solanum chilense, S. peruvianum and S. lycopersicum). The relationships between these traits were tested using linear regression and PCA. There were significant differences in anatomical traits between the species under both OW and RW, where leaf area correlated with stem diameter. Proline and total carbohydrates accumulated highly in S. chilense and S. peruvianum, respectively, and these osmolytes were strongly correlated with increased osmotic potential. Stomatal density varied between species but not between acclimation treatments, while stomatal rate was significantly higher in wild tomatoes. There was a strong positive relationship between stem growth rate and a group of traits together expressed as total stomatal number. Total stomata is described by integration of leaf area, stomatal density, height and internode length. It is proposed that constitutive adaptations and modifications through acclimation that mediate RW play an important role in tolerance to drought stress in herbaceous plants. The capacity for growth under drought stress was not associated with any single combination of traits in wild tomatoes, since the two species differed in relative levels of expression of various phenotypic traits.     

生境异质性及源株密度对互花米草入侵力的影响

外来物种互花米草(Spartina alterniflora)被引种到我国海岸带后,已带来一系列生态危害,严重影响了入侵地生态系统结构和功能,深入了解其入侵机制是实施有效控制的基础和理论依据。长江口的滩涂湿地是我国典型的异质潮滩,拥有多种生境。为了探究异质潮滩环境中互花米草初始源株对其入侵力的影响,研究通过野外移栽实验,模拟不同源株密度,探讨潮滩湿地生境异质性及源株密度对互花米草定居、生长和扩散的影响。结果表明:1)生境异质性及源株密度均对互花米草存活率、株高及种群增长率有极显著影响(P0.01)。2)至生长季末,植被区内互花米草存活率、株高均显著高于前沿光滩(P0.05)。此外,淤泥质潮滩植被区内互花米草株高、种群增长率也显著高于粉砂质潮滩植被区(P0.05)。3)互花米草入侵植被区不受源株密度的限制,但随源株密度增加,互花米草种群增长率显著降低(P0.05),而株高无显著差异(P0.05);在光滩区,只有高源株密度才可入侵成功。4)异质生境的冲淤动态、淹水时间和淹水深度是影响互花米草存活和生长的主要环境因子。丰富了河口湿地互花米草入侵力的机理研究,也为我国开展互花米草防控及海岸带保护、修复和管理提供了科学依据,具有重要的理论和实践意义。     

Associations between macrobenthos and invasive cordgrass, Spartina anglica, in the Danish Wadden Sea

We conducted a survey to provide knowledge on mechanisms controlling spatial and temporal variability of macrobenthos in an intertidal Wadden Sea area partly covered by invasive Spartina anglica. Benthic macrofauna was collected seasonally at seven stations along a transect covering non-vegetated mudflat and vegetated marsh areas. Shannon diversity index was consistently higher for macrobenthos in the open mudflat compared to the marsh area. Infaunal species, like Arenicola marina, Tubificoides benedeni and Macoma balthica, were more abundant in the mudflat than the marsh, while the opposite was evident for epifaunal species, like Hydrobia ulvae and Littorina littorea. The infaunal crustacean, Corophium volutator, on the other hand, appeared particularly attracted to the mudflat–marsh boundary. The biomass of below-ground plant materials and macrodetritus was positively correlated with the total macrofaunal abundance, while the biomass of dead below-ground plant materials alone showed a significant negative correlation with infaunal abundance. Total macrofaunal abundance was highest during summer in the mudflat, while no seasonal pattern was evident in the marsh. The negative influence of S. anglica invasions on infaunal abundance may have serious implications for higher trophic levels, such as waterbirds, that forage in Wadden Sea areas and thus for the overall biodiversity and ecosystem functioning.     

Differentiation and adaptation in <Emphasis Type="Italic">Brassica nigra</Emphasis> populations: interactions with related herbivores

Local adaptation and population differentiation of plants are well documented, but studies on interactions with natural enemies are rare. In particular, evidence for plant adaptation to the local biotic environment, such as herbivores remains poor. We used the black mustard Brassica nigra, an annual species of river valley and coastal habitats to (1) analyse population differentiation in plant traits and herbivory in a common garden experiment, (2) examine home versus away differences in a reciprocal transplant experiment and (3) test whether plants are adapted to local herbivores or vice versa under standard greenhouse conditions. In the common garden experiment, we found significant differentiation in plant traits, leaf damage and herbivore number among seven populations of B. nigra from France and Germany (distance 15–1,000 km). Differences were particularly strong among coastal and river valley populations and did not necessarily increase with geographical distance. A herbivore removal treatment did not change population differentiation when compared with the control allowing natural colonisation. The reciprocal transplant experiment at a scale of 15–30 km did not reveal local plant adaptation, whilst one dominant herbivore species (Meligethes aeneus) occurred in significantly higher numbers on local plants. A greenhouse experiment combining three aphid (Brevicoryne brassicae) and plant populations of the same provenance indicated herbivore adaptation to their local plants rather than plant adaptation, but overall contrasts between local and non-local combinations were not significant. The results suggest that herbivores may counteract local plant adaptation to other environmental factors. Our study has important implications for plant translocations in ecological restoration projects.     

Sexually differential tolerance to water deficiency of Salix paraplesia—A female‐biased alpine willow

Salicaceae plants are dioecious woody plants. Previous studies have shown that male individuals are more tolerant to water deficiency than females for male‐biased poplars. However, Salix paraplesia is a female‐biased species in nature. It is still unknown whether female willows are more tolerant to drought stress than males. To better understand the sexually different tolerance to water deficiency in willows, a greenhouse experiment combined with a field investigation was conducted, and physiological traits were tested in male and female S. paraplesia under a drought‐stressed condition (50% of soil water capacity). Our field investigation showed that S. paraplesia was a species with female‐biased sex ratio along altitude gradients (2,400 m, 2,600 m and 2,800 m) in their natural habitats. Our results showed that the height growth, biomass accumulation, total chlorophyll pigment content (TChl), and the net photosynthetic rate were higher in female willows than in males at the low and middle altitudes (2,400 m and 2,600 m) rather than at a high altitude (2,800 m) under well‐watered conditions. Under drought‐stressed conditions, the growth, biomass, and photosynthesis were greatly inhibited in both sexes, while females showed higher biomass and TChl content and suffered less negative effects than did males. Particularly, females that originated from a high altitude showed lower leaf relative electrolyte leakage, malondialdehyde content, and less disorder of chloroplast ultrastructures but a higher peroxidase activity (POD) than that of males. Therefore, S. paraplesia females exhibited a better drought tolerance and self‐protective ability than males from high altitude. There is a reason to speculate that the population structure of S. paraplesia at a high altitude would be likely to further female biases with the increased drought intensity in the alpine regions.     

Contrasting responses of steppe Stipa ssp. to warming and precipitation variability

Climate change, characterized by warming and precipitation variability, restricted the growth of plants in arid and semiarid areas, and various functional traits are impacted differently. Comparing responses of functional traits to warming and precipitation variability and determining critical water threshold of dominate steppe grasses from Inner Mongolia facilitates the identification and monitoring of water stress effects. A combination of warming (ambient temperature, +1.5°C and +2.0°C) and varying precipitation (?30%, ?15%, ambient, +15%, and +30%) manipulation experiments were performed on four Stipa species (S. baicalensis, S. bungeana, S. grandis, and S. breviflora) from Inner Mongolia steppe. The results showed that the functional traits of the four grasses differed in their responses to precipitation, but they shared common sensitive traits (root/shoot ratio, R/S, and specific leaf area; SLA) under ambient temperature condition. Warming increased the response of the four grasses to changing precipitation, and these differences in functional traits resulted in changes to their total biomass, with leaf area, SLA, and R/S making the largest contributions. Critical water thresholds of the four grasses were identified, and warming led to their higher optimum precipitation requirements. The four steppe grasses were able to adapt better to mild drought (summer precipitation decreased by 12%–28%) when warming 1.5°C rather than 2.0°C. These results indicated that if the Paris Agreement to limit global warming to 1.5°C will be accomplished, this will increase the probability for sustained viability of the Stipa steppes in the next 50–100 years.