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The aim of this study was to explore ecological adaptation of seed microsculptures of Saussurea from different altitudes. Scanning electron microscopy was used to observe the cypselae surface of 10 taxa of Saussurea collected from the eastern edge of Qinghai-Tibet plateau, China. The results indicated that the microsculptures had clear and consistent variations in some samples collected from high (4200 m a.s.l.) and low-altitude sites (2200 m a.s.l.). The thickness of stripes and distance between stripes are increasing while sulcus depth is increasing from a low to a high altitude (P < 0.01). The seed-coat surface was becoming rougher from low to high altitude. The surviving rate of seeds was lower at high altitude than that at low altitude in all the species. The surviving rate of the seeds collected from the high altitudes was higher than that of the seeds collected from low altitudes when sown at the same altitude. All the results suggested that there is selection pressure of the altitude on the development of the microsculpture patterns, which increases absorbed sunlight, the residence time of soil water and nutrition. The changes can make the seeds survive better at extreme environments (cold and dry). The cypselae microsculpture patterns should be used as potential adaptation biomarkers for the species of Saussurea from low- to high-altitude.
Distribution pattern of δ¹³C values of plateau plants and their responses to environment along altitudinal gradients were investigated. In the growing season of 2003 (June– August), stable carbon isotope ratios (δ¹³C) of 174 plant samples belonging to 89 species of 20 families and 58 genera along the gradient 2800– 4400 m (above sea level) was studied in six sites on the east edge of Qinghai-Tibet Plateau. The results indicated that the range of δ¹³C values of plants is narrow from –30.2‰ to –25.2‰, which means that none of the species examined belonged to C₄ photosynthetic pathway and all of these species performed C₃ photosynthetic pathway. The average δ¹³C values of plants at 6 sites were positively correlated to altitude (r = 0.974, P <0.01). The results revealed that site-averaged δ¹³C values were negatively correlated with temperature (r = 0.907, P <0.05) as well as CO₂ partial pressure (r = 0.940, P <0.01). The combination of these two factors account for 80% of the variation of δ¹³C values (r² = 0.859, P <0.01). Varying precipitation with increasing altitude does not affect the plant δ¹³C values (r = 0.469, P> 0.05) as well as the sunlight duration(r = 0.630, P> 0.05).
The frequently changed temperature could have great effects on soil fauna community during soil thawing period in cold areas. Therefore, soil faunas were investigated in both the soil organic layer (OL) and mineral soil layer (ML) in the primary fir (Abies faxoniana) forest (PF), fir and birch (Betula albosinensis) mixed forest (MF) and secondary fir forest (SF) in the eastern Qinghai–Tibet Plateau every ten days between March 5 and April 25, 2009. Soil macrofauna was picked up by hand in the fields. Mesofauna was collected and separated from the soil samples by Baermann and Tullgren methods, respectively. The dominated species of macrofauna belongedto Coleoptera and Diptera at the early stage of soil thawing, and to Coleoptera, Diptera, Araneae and Hymenoptera at the later stage. However, the dominated species of mesofauna belonged to Nematode, Collembola, and Acari in the whole soil thawing. The density, number of taxa, and diversity index of soil fauna showed significant change with temperature fluctuations and reached an obvious peak when soil temperatures rising above 0°C. The density and number of taxa of macrofauna in both the OL and ML were the lowest on March 5 in the three forests, but the density of mesofauna in both the OL and ML was the highest on March 25, except for the ML in the PF. These results implied that soil fauna community was sensitive to temperature fluctuations, which is important in understanding the ecological processes in the winter–spring transitional period.
Because of cold weather and extremely harsh environment, there is few exotic species in Qinghai-Tibetan Plateau. Plateau zokor (Myospalax fontanierii), dominant borrowing small mammal, plays a role as ‘ecosystem engineer’ in alpine meadow ecosystem. We measured and compared the dispersion area, branches, height and number of branches of flixweed tansymustard (Descurainia sophia) on the area disturbed by zokor mounds and in the undisturbed alpine meadow. Flixweed tansymustard is cool-season annual or biennial typical farmland weed in China, and is not found in alpine meadow before. The results indicated that zokor mounds significantly increased the dispersion area, number of individuals in each dispersion area, height and number of branches of flixweed tansymustard on the area disturbed by zokor mounds compared to those in undisturbed alpine meadow. These results suggest that ecosystem engineering by native species of rodent could promote the invasions of alien plant species in alpine meadow ecosystem, leading to higher abundances of invaders.
Given the finite resources of seed production, a trade-off between seed weight and number was expected. However, it still remains unclear how the seed weight-number trade-off changes if there are some mechanisms to enhance resource acquirements during the fruit developments. Sinopodophyllum hexandrum presents furled leaves in flowering time but unfurled leaves in fruiting time, which is considered to enhance the photosynthates to fruit development. To determine the relationship between seed weight and number of S. hexandrum, we examined the altitudinal variations and the relationship between seed weight and number in the fruits from 27 populations of in 2009 and 2010. Our results showed that, along the altitudinal gradient, seed weight decreased significantly but seed number increased insignificantly, indicating that the selective pressure on seed production might come from seed dispersal in high altitude populations. There was no significant relationship between seed weight and number in most populations, but an inverse relationship was found between seed weight and number when the data from all populations were pooled, indicating the seed weight-number trade-off and resource limitation of seed production in S. hexandrum on species level. We suggested that the two unfurled leaves of S. hexandrum might contribute to the resources allocated to the fruit development via enhancing photosynthates, which might mask the seed weight-number trade-off on population level to some degree.
Theoretical and field studies on seed size and plant abundance relationship have been conducted in various communities. However, inconsistent patterns have emerged from these studies, and still little is known about alpine meadows. Here we identified four models and their predictions: the seed size/number trade-off model (SSNTM), the succession model (SM), the spatial competition model (SCM), and the triangle model (TM), in order to assess the relationship between seed size and abundance in alpine meadows, and to elucidate underlying mechanisms. The study site was situated on the eastern Qinghai-Tibetan Plateau at 3500 m above sea level. From 1999 through 2001, two indices of plant abundance (aboveground biomass and density) were simultaneously measured in 45 quadrates (0.25 m²). Data for 101 plant species (mostly Cyperaceae, Poaceae, Asteraceae, Ranunculaceae and forbs) showed that seed size is like log normal distributed, and it slightly skewed in smaller-sized seeds. The SSNTM, the TM, the SM and the SCM models were not supported in this alpine meadow, and the relationship between seed size and abundance was always positive (although in some samples, the relationship was not significant). The positive correlation between seed size and abundance observed for some grassland communities was also demonstrated in the alpine meadow. It suggests that seed size depends on the plant growth form, but the biomass-density relationship is inconsistent with previous studies. This suggests that the measure of abundance used in these studies is not the only reason for inconsistency of seed size.
It has been observed that leaf morphology shift within species is linked to climate change, but there are few studies on the effects of altitude change on leaf morphology of species. We hypothesized that similar to climate change, a morphological shift within species would occur over time under different growing altitudes. In this study, we evaluated three dominant grass species: Elymus nutans Griseb., Kobresia capillifolia Clarke., Carex moorcroftii Boott., taking advantage of the altitudinal variations (3000-4000 a.s.l.) on the Qinghai-Tibetan Plateau. Our study showed that almost all leaf traits of these three species had significant differences (P <0.05) across an altitudinal gradient. Different species responded differently to altitude change. Leaf thickness (LT) of the three species increased with increase in altitude. Leaf area (LA) of E. nutans and C. moorcroftii decreased with increasing altitude, but that of K. capillifolia increased. There was no obvious linear effect on leaf dry matter content (LDMC) and specific leaf area (SLA) of these three species. LDMC of E. nutans and C. moorcroftii showed a trend of increase, while that of K. capillifolia decreased. SLA of E. nutans and K. capillifolia showed a trend of increase, but that of C. moorcroftii decreased with increase in altitude. In addition, soil pH (pH) and air temperature (AT) decreased with increase in altitude. However, other soil and climate factors increased as altitude increased. The finding of this work is that leaf morphology shift within species happens under altitude change to adapt to specific environment.
Subterranean life style is characterized by limited food resource and energy consuming burrow behavior. Subterranean rodents are usually recognized as dietary generalists. In the current study, we investigated the species composition of winter caches of plateau zokors (Eospalax baileyi), and the dissimilarity of species composition in caches and in the vicinity of the burrows. We have made our research in Kobresia dominated vegetation at the altitude 3200 m a.s.l. We used the non-parametric method to test the similarities between the relative dry mass of the plant species within caches and their closed vicinity. This method was based on measures of distance between pairs of individual multivariate observations. Our results revealed that although plateau zokors cached most (60) of the plant species found in the vicinity of their burrow systems (66), the cached dry mass was dominated by a few species. Nine plant species accounted for 80% dry mass of plants present in zokors’ caches, Polygonum viviparum (50%) and other perennial poisonous forbs constitute 70% dry mass of the caches. Results showed that the dissimilarity between the relative dry mass of plant species within caches and in their vicinity was extremely significant (for Bray-Curtis distance measure, R = 0.355, P = 0.000; for Euclidean distance measure R =0.319, P =0.000). These results showed that plateau zokors don’t forage randomly when collecting food for winter caches. We suppose that rather than using a non-selective foraging strategy by dealing with high fiber-content plants, plateau zokors have forged another dietary strategy to meet the energy demands in the evolutionary process: to deal with poisonous forbs.
The ongoing warming in the Qinghai-Tibetan Plateau leads to changes in ecosystem processes while the responses of soil and vegetation are not well understand. Thus, we used infrared radiators to carry out experimental warming from July 2010 to August 2011 in an alpine meadow on the Plateau (about 4630 m above sea level) to research the responses of environmental factors and vegetation characteristics to short-term warming (1 year). The experimental design was a block design consisting of five replications and included three treatment levels: control, T1 (130 W m–2) and T2 (150 W m–2). The results showed that air temperature at 20 cm height, surface temperature and soil temperature in the 0–100 cm layers increased with warming. The biggest differences of T1 (1.66°C) and T2 (2.34 °C) appeared on the surface and at 20 cm depth, whereas the biggest amplitudes of T1 (27.15%) and T2 (35.81%) all occurred at 100 cm depth. Soil moisture showed different trends with warming in different soil layers. In the 0–40 cm layers, soil moisture decreased with warming. The biggest differences (–2.97% for T1 and –2.73% for T2) and amplitudes (–18.07% for T1 and –16.64% for T2) all appeared at 10 cm depth. In the 60–100 cm layers, soil moisture increased with warming. The biggest differences (2.53% for T1 and 6.45% for T2) and amplitudes (11.39% for T1 and 29.05% for T2) all occurred at 100 cm depth. Relative to control, vegetation height and aboveground biomass increased significantly in T1 and T2 (P <0.05), while vegetation coverage had not significant differences in T1 and T2 (P> 0.05). In T1 and T2, the amplitudes were 30.67% and 30.19% for vegetation height, and 36.22% and 27.87% for vegetation aboveground biomass, and 12.89% and 4.42% for vegetation coverage, respectively. In the path analysis between environment and vegetation properties, vegetation was directly affected by soil moisture at 40 cm and 60 cm depths, whereas indirectly influenced by relative humidity at 20 cm height and soil temperature at 40 cm depth. This might be related to the downward movement of the soil moisture caused by warming.
The influence of air and soil warming on root vole (Microtus oeconomus L.) population was studied in winter period in top open chambers (OTC) (0.8–1.8 m²) warmed by conical fiberglass material and situated in alpine meadow (3250 m) at Qinghai-Tibet Plateau, China. The OTCs were distributed on an area of 30 × 30 m of experimental warming site; another site of the same area was a control one. The root vole population was investigated on two pairs of sites in “low-grazing” and “high-grazing” (by sheep) parts of the meadow; mark-recapture method was used. The winter-season averaged air and soil temperature inside of the chambers were 1.3°C higher than the temperature outside the chambers. The warming in the chambers had no statistically significant effect on root vole numbers, on average body mass of individual, and on average body mass of males and females. In conclusion, as small as 1.3°C warming of soil and air introduced locally and on small (several m²) scale, in the alpine meadow habitat in winter period, has possibly no effect on root vole numbers and biomass.
Summer diets of two sympatric raptors Upland Buzzards (Buteo hemilasius Temminck et Schlegel) and Eurasian Eagle Owls (Bubo bubo L. subsp. Hemachalana Hume) were studied in an alpine meadow (3250 m a.s.l.) on Qinghai- Tibet Plateau, China. Root voles Microtus oeconomus Pallas, plateau pikas Ochotona curzoniae Hodgson, Gansu pikas O. cansus Lyon and plateau zokors Myospalax baileyi Thomas were the main diet components of Upland Buzzards as identified through the pellets analysis with the frequency of 57, 20, 19 and 4%, respectively. The four rodent species also were the main diet components of Eurasian Eagle Owls basing on the pellets and prey leftovers analysis with the frequency of 53, 26, 13 and 5%, respectively. The food niche breadth indexes of Upland Buzzards and Eurasian Eagle Owls were 1.60 and 1.77 respectively (higher value of the index means the food niche of the raptor is broader), and the diet overlap index of the two raptors was larger (Cue = 0.90) (the index range from 0 – no overlap – to 1 – complete overlap). It means that the diets of Upland Buzzards and Eurasian Eagle Owls were similar (Two Related Samples Test, Z = –0.752, P = 0.452). The classical resource partitioning theory can not explain the coexistence of Upland Buzzards and Eurasian Eagle Owls in alpine meadows of Qinghai-Tibet Plateau. However, differences in body size, predation mode and activity rhythm between Upland Buzzards and Eurasian Eagle Owls may explain the coexistence of these two sympatric raptors.
Grazing can change plant community composition and structure, which may alter the functions of the shrub meadow ecosystem. Grazing effects on Potentilla fruticosa shrub community in the headwater region of the Yellow River, which is in core area of the Qinghai-Tibet Plateau, are studied to provide adequate protection decision-making. We investigated continuous grazing and seasonal enclosure effects on P. fruticosa shrub communities. Three sites of P. fruticosa shrub comprising both continuous grazing and seasonal enclosure treatments were selected. The size of each fenced plot of P. fruticosa shrub was about 3000 m², the stocking rate was about 5 heads per 100 m² in continuous grazing treatment. Three samplings were made in each growing season of 2003 and 2004. Cover of vegetation, plant species composition and vegetation height were investigated in seven 1 × 1 m quadrates in each treatment. Above-ground biomass was measured in five 0.5 × 0.5 m quadrates. Shrub, forb, graminoid and sedge plant materials were clipped at ground level and oven-dried at 85ºC to a constant mass. Plant composition was affected by long term continuous grazing and changes were caused by forb species shifting. No apparent difference in species richness between the grazed and ungrazed communities over the growing months were found but the Shannon’s diversity indices of the grazed communities in June and July were higher than that of the ungrazed but lower in the late August and September. Live vegetation cover was reduced by 6.7%, 7.3% and 11.5%, respectively, owing to grazing in July, August and September, but not in June (P> 0.05). Forbs took up more than 50% cover of the vegetation in both grazed and ungrazed treatments. Relative cover of sedges and forbs in ungrazed treatment decreased in July, August and September, while that of graminoids increased more than 70% in the same period. Live vegetation height was reduced by 27% (2004) and 23% (2003) in late August and early September, but not in early growing season. Grazing reduced total above-ground biomass by 35%, 37% and 36% in July, August and early September, respectively, and the reduction was mainly in forb biomass. Continuous grazing affects plant composition and species diversity. The quantitative characteristics of P. fruticosa communities were influenced by grazing over growing months, but the effects were offset by non-growing season grazing.
Mitochondrial cytochrome b gene sequences and morphological data (body length, hind foot length, etc.) for twelve populations with pairwise distances 27–600 km in Qinghai-Tibetan Plateau (distributed in Qinghai, Gansu and Sichuan Province, and at the altitude 3020–4550 m) in Western China were used to investigate the phylogeographical pattern of Plateau zokor (Myospalax baileyi Thomas). There was a little disparity between mtDNA genetic distance and morphological Euclidean distance on population relationships. However, there is a significant correlation (P <0.001) calculated by Mantel’s tests was validated between mtDNA and morphology distances. Analysis of Molecular Variance showed that most of the observed genetic variations occurred between populations, indicating little maternal gene flow between them, as a result of geographical restrictions. Phylogenetic analysis coupled with cluster analysis together showed that the substantial population structuring and phylogenetic discontinuities existed within this species. The evident allopatric population structuring of this subterranean rodent may mostly result from its specialized subterranean excavating behavior with high energy costs, predation from grassland raptors and also the influences of perennial tundra and environmental desiccation in the Qinghai-Tibetan Plateau.
Long-term overgrazing has resulted in grassland deterioration and even desertification on the eastern Qinghai-Tibetan Plateau. In this paper, we examined the characteristics of vegetation and soil properties in the livestock-excluded pastures and the adjacent grazed pastures under two topographic habitats (the flat valley and the south-facing slope). Seven-year exclusion of livestock has enhanced aboveground live biomass, root biomass and litter accumulation. Livestock exclusion has also increased soil bulk density and soil water content, soil organic C concentration, total N concentration and its transformation rate, and soil microbial activity. The results showed that livestock exclusion has facilitated vegetation recovery and improved physical, chemical and biological properties of soil. However, livestock exclusion has significantly decreased graminoid biomass accumulation, especially on the flat valley, the biodiversity also significantly decreased there. The results suggested that long-term livestock exclusion was disadvantageous for palatable forage production and biodiversity protection on the flat valley. Compared to the flat valley, the grassland on the south-facing slope was under more severe degradation, and the reversion was in a slower process. Thus, the optimal grassland management in the livestock-excluded pasture on the flat valley should include a low or moderate grazing intensity or adopt an alternate grazing system, but more effective and even longer livestock exclusion practice should be taken on the south-facing slope.
It is known that the dominance of graminoid species is promoted by N addition; however, there has been relatively little effort to examine the pattern induced by natural N fertility. Since nutrie nt use e fficiency (NUE) is an important trait determining plant competitive ability, we expected that the species guild with higher NUE (lower nutrient content) may be more competitive on infertile soils. We explo red t he relationships between relative forbs biomass share , soil N and productivity by the linear regressi on analysis on a natural alpine meadow in northeast of Qinghai- Tibetan Plateau (3600 m a.s.l.). To test the variety of leaf N:P stoichiometry, paired t test and general linear model multivariate (GLM) analysis were also used. We f ound that the leaf N:P ratios of the whole community were below 13 in studied sites, which may be consistent with the N limitation on the veget ation. Graminoids re tained lower concentrations of leaf N and P than forbs in community on the Nlimited grassland. Consistent with our prediction, we found that the biomass fraction of graminoids declined with soil N content and aboveground production on the grassland. Different from the pattern along fertility gradients induced by N fertilization, our results showed that gr aminoids with lower internal nutrient content w ere able to resist low levels of nutrient availability on the natural alpine grassland when compared to forbs.
Grassland degradation due to anthropogenic and natural factors and their interactions is one of the worldwide ecological and economic problems because it reduces grassland productivity and diversity and leads to desertification. The objective of this study was to assess the influence of protective enclosure on vegetation composition and diversity and plant biomass of an alpine degraded meadow. The experiment was conducted at center of Qinghai-Tibetan Plateau with two degraded (caused by overgrazing) alpine meadows: the lightly and severely degraded ones (LD and SD) and their enclosed areas with iron net (LDE and SDE, respectively). The areas 200 m × 150 m for each of four degraded alpine meadow treatments at average altitude 3,960 m a.s.l. were set for research. The lightly degraded plots were dominated by Scirpus distigmaticus (Kukenth.) Tang et Wang, Elymus nutans Griseb. and Oxytropis ochrocephala Bunge. The dominating plants in severely degraded plots were: Artemisia sieversiana Ehrhart ex Willd, Ajania tenuifolia (Jacq.) Tzvel, Lonicera minuta Batal. The results showed: (1) the vegetation cover of two degraded plots (LD and SD) has increased after taking the enclosure measures and the forbs dominated both plots. (2) Species richness has also increased in two enclosed degraded plots, respectively. There no significant differences in evenness and diversity between LD and LDE, and SD and SDE, respectively. (3) Enclosure may promote aboveground biomass, particularly grass and forb biomass in LD, and forb biomass in SD plots.
In this study, somatic chromosome counts were determined in 302 individuals from 43 Allium przewalskianum populations; 90 were diploids (2n = 16, 16 + 1B, 16 + 2B) and 212 were tetraploids (2n = 32). Of the 43 populations, five were selected for karyotype analysis. Among them, the diploid plants have two karyotypes: 2n = 2x = 14m + 2st (2SAT) and 2n =2x = 8m + 6sm + 2st (2SAT). The tetraploid has one: 2n = 4x = 28m + 4st. Mixoploidy, Robertsonian translations, and B chromosomes were reported for the first time. In combination with previous chromosome data, the present study reveals a uniform basic chromosomal number (x = 8) and uniformity of karyotypes (Stebbins's 2A type), indicating that speciation through polyploidization is less likely in A. przewalskianum, despite the highly diversified habitats it occupies.
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