No C_4 Plants Found at the Haibei Alpine Meadow Ecosystem Research Station in Qinghai, China: Evidence from Stable Carbon Isotope Studies

Using the measurement of stable carbon isotopes in leaves as a tool to investigate photosyn-thetic pathway of 102 plant species grown at an alpine meadow ecosystem, at the foot of the Qilian Mountain, Qinghai, China. The results indicate that the δ~(3)C values of plants have a narrow range from -28.24‰ to -24.84‰, which means that none of the species examined belongs to C_4 and crassulaceous acid metabolism (CAM) photosynthetic pathway and all of these species perform photosynthesis through the C_3 pathway. This is likely due to a long-term adaptation to environments at the alpine meadow ecosystem.

Effect of plateau zokors on vegetation characteristics and productivity of alpine meadow

This research was conducted on alpine meadow site at Menyuan county, Qinghai Province, People's Republic of China to determine the effects of native, subterranean rodent of Qinghai-Tibet grasslands, the plateau zokors (Myospalax baileyi), on seasonal above-and below-ground plant biomass, plant species diversity and productivity. Both total peaks of above-and below-ground biomass were the greatest (413.600 g/m~2 and 2297.502 g/m~2) in the patch no any plateau zokors colonized by plateau zokors over 10 years in August and October, respectively. Both above-and below-ground biomass were significantly increased in the patches where plateau zokors were removed or the burrow systems were abandoned for five years compared to the patches plateau zokors colonized over 10 years. However, both above-and below-ground biomass in abandoned patches were significantly lower than that in uncolonized patches. Monocotyledonous biomass was reduced greatly, but the non-palatable dicots were significantly increased in colonized patches. The palatable biomass of monocots and dicots were increased in abandoned patches. Total plant species diversity was the greatest in uncolonized patchesand least in abandoned patch. The total net primary production in colonized patches was reduced by 68.98% compared with uncolonized patches. Although the patches were without any plateau zokors disturbance for fives years, the total net primary production just reached 58.69% of the uncolonized patches. The above-ground net primary production in abandoned patches increased 28.74% and the below-ground increased 54.91% compared with the colonized patches. We suggest that plateau zokor-induced changes in plant above- and below-ground biomass and species diversity may lead to further alterations of nutrient cycling and trophic dynamics in this alpine meadow ecosystem.

Methane emissions by alpine plant communities in the Qinghai-Tibet Plateau

For the first time to our knowledge, we report here methane emissions by plant communities in alpine ecosystems in the Qinghai-Tibet Plateau. This has been achieved through long-term field observations from June 2003 to July 2006 using a closed chamber technique. Strong methane emission at the rate of 26.2 +/- 1.2 and 7.8 +/- 1.1 mu g CH4 m(-2) h(-1) was observed for a grass community in a Kobresia humilis meadow and a Potentilla fruticosa meadow, respectively. A shrub community in the Potentilla meadow consumed atmospheric methane at the rate of 5.8 +/- 1.3 mu g CH4 m(-2) h(-1) on a regional basis; plants from alpine meadows contribute at least 0.13 Tg CH4 yr(-1) in the Tibetan Plateau. This finding has important implications with regard to the regional methane budget and species-level difference should be considered when assessing methane emissions by plants.

Nitrous oxide emissions from two alpine meadows in the Qinghai-Tibetan Plateau

Nitrous oxide (N2O) emission was measured in a Kobresia humilis meadow and a Potentilla fruticosa meadow in the Qinghai-Tibet Plateau from June 2003 to July 2006. Five treatments were setup in the two alpine meadows. Two bare soil treatments were setup in the K. humilis meadow (BSK) and in the P. fruticosa meadow (BSP) by removing the above- and belowground plant biomass. Three plant community treatments were setup with one in the K. humilis meadow (herbaceous community in the K. humilis meadow-HCK) and two in the P. fruticosa meadow (herbaceous community in the P. fruticosa meadow-HCP, and shrub community in the P. fruticosa meadow-SCP). Nitrous oxide emission from BSP was estimated to be 38.1 +/- 3.6 mu g m(-2) h(-1), significantly higher than from BSK (30.2 +/- 2.8 mu g m(-2) h(-1)) during the whole experiment period. Rates from the two herbaceous blocks (HCK and HCP) were close to 39.5 mu g m(-2) stop h(-1) during the whole experimental period whereas shrub community (SCP) showed significant high emission rates of N2O. Annual rate of N2O emission was estimated to be 356.7 +/- 8.3 and 295.0 +/- 11.6 mg m(-2) year(-1) from the alpine P. fruticosa meadow and from the alpine K. humilis meadow, respectively. These results suggest that alpine meadows in the Qinghai-Tibetan Plateau are an important source of N2O, contributing an average of 0.3 Tg N2O year(-1). We concluded that N2O emission will decrease, due to a predicted vegetation shift from shrubs to grasses imposed by overgrazing.

Small-scale species richness and its spatial variation in an alpine meadow on the Qinghai-Tibet Plateau

We investigated how the high small-scale species richness of an alpine meadow on the Qinghai-Tibet Plateau, China, is maintained. This area is characterized by strong wind and severe cold during long winters. In winter, most livestock is grazed on dead leaves in small pastures near farmers' residences, whereas in the short summer, livestock is grazed in mountainous areas far from farmers' residences. The number of plant species and the aboveground biomass were surveyed for three adjacent pastures differing in grazing management: a late-winter grazing pasture grazed moderately from 1 February to 30 April, an early-winter grazing pasture grazed lightly from 20 September to late October, and a whole-year grazing pasture grazed intensively throughout the entire year. In each pasture, we harvested the aboveground biomass from 80 or 100 quadrats of 0.01 m(2) along a transect and classified the contents by species. We observed 15.5-19.7 species per 0.01 m(2), which is high richness per 0.01 m(2) on a worldwide scale. The species richness in the two winter grazing pastures was higher than that in the whole-year grazing pasture. The spatial variation in species richness and species composition in the two winter grazing pastures in which species richness was high was greater than that in the whole-year grazing pasture in which species richness was lower. Most of the leaves that are preserved on the winter grazing pastures during summer are blown away by strong winds during winter, and the remaining leaves are completely exhausted in winter by livestock grazing. A pasture with a high richess is accompanied by a high spatial variation in species richness and species composition. There is a high possibility that the characteristic of spatial variation is also caused by traditional grazing practices in this area.

Response of Polygonum viviparum species and community level to long-term livestock grazing in alpine shrub meadow in Qinghai-Tibet Plateau

Grazing by domestic herbivores is generally recognized as a major ecological factor and an important evolutionary force in grasslands. Grazing has both extensive and profound effects on individual plants and communities. We investigated the response patterns of Polygonum viviparum species and the species diversity of an alpine shrub meadow in response to long-term livestock grazing by a field manipulative experiment controlling livestock numbers on the Qinghai-Tibet Plateau in China. Here, we hypothesize that within a range of grazing pressure, grazing can alter relative allocation to different plant parts without changing total biomass for some plant species if there is life history trade-offs between plant traits. The same type of communities exposed to different grazing pressures may only alter relative species' abundances or species composition and not vary species diversity because plant species differ in resistant capability to herbivory. The results show that plant height and biomass of different organs differed among grazing treatments but total biomass remained constant. Biomass allocation and absolute investments to both reproduction and growth decreased and to belowground storage increased with increased grazing pressure, indicating the increasing in storage function was attained at a cost of reducing reproduction of bulbils and represented an optimal allocation and an adaptive response of the species to long-term aboveground damage. Moreover, our results showed multiform response types for either species groups or single species along the gradient of grazing intensity. Heavy grazing caused a 13.2% increase in species richness. There was difference in species composition of about 18%-20% among grazing treatment. Shannon-Wiener (H') diversity index and species evenness (E) index did not differ among grazing treatments. These results support our hypothesis.

Feeding behaviour of yaks on spring, transitional, summer and winter pasture in the alpine region of the Qinghai-Tibetan plateau

Of the present estimated world population of 14.2 million yaks, approximately 13.3 million occur within Chinese territories (Food and Agriculture Organization of the United Nations, 2003). Although there is an extensive bibliography covering the species, few studies have been conducted in the area of foraging behaviour. The present study was conducted at pasture during the spring, transitional, summer and winter seasons to determine the daily temporal patterns of grazing and ruminating behaviour by yaks. During each study period, two 24 h recordings were undertaken with each of six mature dairy yaks. One study period was conducted on each of the transitional, summer and winter pastures, whereas, due to the considerable changes occurring in the morphology of the spring pasture, three separate studies were completed during March, April and May. During the second of these studies (April), the effect of level of concentrate supplementation on grazing and ruminating behaviour was also examined. Behaviour recordings were made using solid-state behaviour recorders. Short-term intake rates (IR, g min(-1)) were calculated by weighing yaks before and after approximately 1 h of grazing, retaining the faeces and urine excreted and applying a correction for insensible weight loss. Yaks spent less time grazing during the dry season (the early period on the spring pasture) compared with the later green swards (the later period on the spring pasture, the transitional pasture and the summer pasture) (P < 0.05). When the forage quality improved, but there was still insufficient mass (the later period on the spring pasture), the yaks extended their grazing time at the expense of other activities. During the early periods on the spring pasture, the short-term IR by yaks was up to 53 g DM min(-1), significantly higher than at other times (P < 0.05). The level of concentrate offered had little or no effect on grazing or ruminating time. The total eating time of the yaks offered 0.5 or 1.0 kg concentrate was 2.9 and 4.5 h day(-1) respectively, significantly lower than unsupplemented yaks (6.8 h) (P < 0.05). In general, yaks can regulate their foraging behaviour according to the changes of sward conditions in order to achieve optimal grazing strategies. (C) 2007 Published by Elsevier B.V.

Potential refugium on the Qinghai-Tibet Plateau revealed by the chloroplast DNA phylogeography of the alpine species Metagentiana striata (Gentianaceae)

Metagentiana striata is an alpine annual herbaceous plant endemic to the east of the Qinghai-Tibet (Q-T) Plateau and adjacent areas. The phylogeography of M. striata was studied by sequencing the chloroplast DNA (cpDNA) trnS-trnG intergenic spacer. Ten haplotypes were identified from an investigation of 232 individuals of M. striata from 14 populations covering the entire geographical range of this species. The level of differentiation amongst populations was very high (G(ST) = 0.746; N-ST = 0.774) and a significant phylogeographical structure was observed (P < 0.05). An analysis of molecular variance found a high variation amongst populations (76%), with F-ST = 0.762 (highly significant, P < 0.001), indicating that little gene flow occurred amongst the different regions; this was explained by the isolation of populations by high mountains along the Q-T Plateau and adjacent areas (N-m = 0.156). Only one ancestral haplotype (A) was common and widespread throughout the distributional range of M. striata. The populations of the Hengduan Mountains region of the south-eastern Q-T Plateau showed high diversity and uniqueness of haplotypes. It is suggested that this region was the potential refugium of M. striata during the Quaternary glaciation, and that interglacial and postglacial range expansion occurred from this refugium. This scenario was in good agreement with the results of nested clade analysis, which inferred that the current spatial distribution of cpDNA haplotypes and populations resulted from range expansion, together with past allopatric fragmentation events. (c) 2008 The Linnean Society of London.

Summer diet of two sympatric species of raptors Upland Buzzard (Buteo hemilasius) and Eurasian Eagle Owl (Bubo bubo) in alpine meadow: problem of coexistence

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 (C-ue = 0.90) (the index range from 0 - no overlap - to I - 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.

Growing season ecosystem respirations and associated component fluxes in two alpine meadows on the Tibetan Plateau

From 30 June to 24 September in 2003 ecosystem respiration (Re) in two alpine meadows on the Tibetan Plateau were measured using static chamber- and gas chromatography- (GC) based techniques. Simultaneously, plant removal treatments were set to partition Re into plant autotrophic respiration (Ra) and microbial heterotrophic respiration (Rh). Results indicated that Re had clear diurnal and seasonal variation patterns in both of the meadows. The seasonal variability of Re at both meadow sites was caused mainly by changes in Ra, rather than Rh. Moreover, at the Kobresia humilis meadow site (K_site), Ra and Rh accounted for 54% and 46% of Re, respectively. While at the Potentilla fruticosa scrub meadow (P_site), the counterparts accounted for 61% and 39%, respectively. T test showed that there was significant difference in Re rates between the two meadows (t = 2.387, P = 0.022). However, no significant difference was found in Rh rates, whereas a significant difference was observed in Ra rates between the two meadows. Thus, the difference in Re rate between the two meadows was mainly attributed to plant autotrophic respirations. During the growing season, the two meadows showed relatively low Q(10) values, suggesting that Re, especially Rh was not sensitive to temperature variation in the growing season. Additionally, Re and Rh at the K_site, as well as Rh at the P_site was negatively correlated with soil moisture, indicating that soil moisture would also play an important role in respirations.

Altitudinal differences in the leaf fitness of juvenile and mature alpine spruce trees (Picea crassifolia)

In many plant species, leaf morphology varies with altitude, an effect that has been attributed to temperature. It remains uncertain whether such a trend applies equally to juvenile and mature trees across altitudinal gradients in semi-arid mountain regions. We examined altitude-related differences in a variety of needle characteristics of juvenile (2-m tall) and mature (5-m tall) alpine spruce (Picea crassifolia Kom.) trees growing at altitudes between 2501 and 3450 m in the Qilian Mountains of northwest China. We found that stable carbon isotope composition (delta C-13), area- and mass-based leaf nitrogen concentration (N-a, N-m), number of stomata per gram of nitrogen (St/N), number of stomata per unit leaf mass (St/LM), projected leaf area per 100 needles (LA) and leaf mass per unit area (LMA) varied nonlinearly with altitude for both juvenile and mature trees, with a relationship reversal point at about 3 100 m. Stomatal density (SD) of juvenile trees remained unchanged with altitude, whereas SD and stomatal number per unit length (SNL) of mature spruce initially increased with altitude, but subsequently decreased. Although several measured indices were generally found to be higher in mature trees than in juvenile trees, N-m, leaf carbon concentration (C.), leaf water concentration. (LWC), St/N, LA and St/LM showed inconsistent differences between trees of different ages along the altitudinal gradient. In both juvenile and mature trees, VC correlated significantly with LMA, N-m, N-a, SNL, St/LM and St/N. Stomatal density, LWC and LA were only significantly correlated with delta C-13 in mature trees. These findings suggest that there are distinct ecophysiological differences between the needles of juvenile and mature trees that determine their response to changes in altitude in semi-arid mountainous regions. Variations in the fitness of forests of different ages may have important implications for modeling forest responses to changes in environmental conditions, such as predicted future temperature increases in high attitude areas associated with climate change.

Cellular automata: Simulating alpine tundra vegetation dynamics in response to global warming

This study attempts to model alpine tundra vegetation dynamics in a tundra region in the Qinghai Province of China in response to global warming. We used Raster-based cellular automata and a Geographic Information System to study the spatial and temporal vegetation dynamics. The cellular automata model is implemented with IDRISI's Multi-Criteria Evaluation functionality to simulate the spatial patterns of vegetation change assuming certain scenarios of global mean temperature increase over time. The Vegetation Dynamic Simulation Model calculates a probability surface for each vegetation type, and then combines all vegetation types into a composite map, determined by the maximum likelihood that each vegetation type should distribute to each raster unit. With scenarios of global temperature increase of I to 3 degrees C, the vegetation types such as Dry Kobresia Meadow and Dry Potentilla Shrub that are adapted to warm and dry conditions tend to become more dominant in the study area.

Changes in plant biomass and species composition of alpine Kobresia meadows along altitudinal gradient on the Qinghai-Tibetan Plateau

Alpine Kobresia meadows are major vegetation types on the Qinghai-Tibetan Plateau. There is growing concern over their relationships among biodiversity, productivity and environments. Despite the importance of species composition, species richness, the type of different growth forms, and plant biomass structure for Kobresia meadow ecosystems, few studies have been focused on the relationship between biomass and environmental gradient in the Kobresia meadow plant communities, particularly in relation to soil moisture and edaphic gradients. We measured the plant species composition, herbaceous litter, aboveground and belowground biomass in three Kobresia meadow plant communities in Haibei Alpine Meadow Ecosystem Research Station from 2001 to 2004. Community differences in plant species composition were reflected in biomass distribution. The total biomass showed a decrease from 13196.96 +/- 719.69 g/m(2) in the sedge-dominated K. tibetica swamp to 2869.58 +/- 147.52 g/m(2) in the forb and sedge dominated K. pygmaea meadow, and to 2153.08 +/- 141.95 g/m(2) in the forbs and grasses dominated K. humilis along with the increase of altitude. The vertical distribution of belowground biomass is distinct in the three meadow communities, and the belowground biomass at the depth of 0-10 cm in K. tibetica swamp meadow was significantly higher than that in K. humilis and K. pygmaea meadows (P < 0.01). The herbaceous litter in K. tibetica swamp was significantly higher than those in K. pygnaeca and K. humilis meadows. The effects of plant litter are enhanced when ground water and soil moisture levels are raised. The relative importance of litter and vegetation may vary with soil water availability. In the K. tibetica swamp, total biomass was negatively correlated to species richness (P < 0.05); aboveground biomass was positively correlated to soil organic matter, soil moisture, and plant cover (P < 0.05); belowground biomass was positively correlated with soil moisture (P < 0.05). However, in the K. pygnaeca and K. humilis meadow communities, aboveground biomass was positively correlated to soil organic matter and soil total nitrogen (P < 0.05). This suggests that the distribution of biomass coincided with soil moisture and edaphic gradient in alpine meadows.

Genetic variation in the endangered Anisodus tanguticus (Solanaceae), an alpine perennial endemic to the Qinghai-Tibetan Plateau

We used random amplified polymorphic DNA markers (RAPDs) to assess genetic variation between- and within-populations of Anisodus tanguticus (Solanaceae), an endangered perennial endemic to the Qinghai-Tibetan Plateau with important medicinal value. We recorded a total of 92 amplified bands, using 12 RAPD primers, 76 of which (P = 82.61%) were polymorphic, and calculated values of H-t and H-sp of 0.3015 and 0.4459, respectively, suggesting a remarkably high rate of genetic variation at the species level. The average within-population diversity also appeared to be high, with P, H-e and H-pop values of 55.11%, 0.1948 and 0.2918, respectively. Analyses of molecular variance (AMOVA) showed that among- and between-population genetic variation accounted for 67.02% and 32.98% of the total genetic variation, respectively. In addition, Nei's coefficient of differentiation (G(ST)) was found to be high (0.35), confirming the relatively high level of genetic differentiation among the populations. These differentiation coefficients are higher than mean corresponding coefficients for outbreeding species, but lower than reported coefficients for some rare species from this region. The genetic structure of A. tanguticus has probably been shaped by its breeding attributes, biogeographic history and human impact due to collection for medicinal purposes. The observed genetic variations suggest that as many populations as possible should be considered in any planned in situ or ex situ conservation programs for this species.

Effect of imitated global warming on Delta C-13 values in seven plant species growing in Tibet alpine meadows

Open-top chambers were used to estimate the possible effects of global warming on delta C-13 of seven plant species grown in alpine meadow ecosystem. The delta C-13 values of plant species were lower after long-term growth in open-top chambers. In the course of experiment, temperature significantly increased inside the chambers by 4 degrees C. Plant species grown at a lower elevation above sea level had higher delta C-13 values as compared to those grown at a higher elevation. This was in accordance with the effect of open-top chamber on delta C-13 values in plants. Greater availability of CO2 and lower water vapor as indicated by an increase in discrimination against (CO2)-C-13, probably result in more negative delta C-13 values of plants because higher stomatal conductance increases availability of CO2 and causes greater discrimination against (CO2)-C-13. The plant species studied could be the indicator species for testing global warming by the change in carbon isotope ratios at the two growth temperatures.