Decline in medicinal and forage species with warming is mediated by plant traits on the Tibetan Plateau

Experimental studies of how global changes and human activities affect plant diversity often focus on broad measures of diversity and discuss the implications of these changes for ecosystem function. We examined how experimental warming and grazing affected species within plant groups of direct importance to Tibetan pastoralists: medicinal plants used by humans and palatable plants consumed by livestock. Warming resulted in species losses from both the medicinal and palatable plant groups; however, differential relative vulnerability to warming occurred. With respect to the percent of warming-induced species losses, the overall plant community lost 27%, medicinal plants lost 21%, and non-medicinal plants lost 40% of species. Losses of palatable and non-palatable species were similar to losses in the overall plant community. The deep-rootedness of medicinal plants resulted in lowered sensitivity to warming, whereas the shallow-rootedness of non-medicinal plants resulted in greater sensitivity to warming; the variable rooting depth of palatable and non-palatable plants resulted in an intermediate response to warming. Predicting the vulnerability of plant groups to human activities can be enhanced by knowledge of plant traits, their response to specific drivers, and their distribution within plant groups. Knowledge of the mechanisms through which a driver operates, and the evolutionary interaction of plants with that driver, will aid predictions. Future steps to protect ecosystem services furnished by medicinal and palatable plants will be required under the novel stress of a warmer climate. Grazing may be an important tool in maintaining some of these services under future warming.

Status and dynamics of Kobresia pygmaea ecosystem on the Tibetan plateau

This paper provides information about the distribution, structure, and ecology of the world's largest alpine ecosystem, the Kobresia pygmaea pastures in the southeastern Tibetan plateau. The environmental importance of these Cyperaceae mats derives from the extremely firm turf, which protects large surfaces against erosion, including the headwaters of the Huang He, Yangtze, Mekong, Salween, and Brahmaputra. The emphasis of the present article is on the climate-driven evolution and recent dynamics of these mats under the grazing impact of small mammals and livestock. Considering pedological analyses, radiocarbon datings, and results from exclosure experiments, we hypothesize that the majority of K. pygmaea mats are human-induced and replace forests, scrub, and taller grasslands. At present, the carrying capacity is increasingly exceeded, and reinforced settlement of nomads threatens this ecosystem especially in its drier part, where small mammals become strong competitors with livestock and the removal of the turf is irreversible. Examples of rehabilitation measures are given.

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.

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.

Competition between Stipa grandis and Cleistogenes squarrosa

Competitive strategy for resources between Cleistogenes squarrosa (Trin.) Keng which is a dominant species of grasslands degraded by moderate-heavy grazing, and Stipa grandis P. Smirnov, which is a dominant species of ungrazed communities, was studied using a replacement series method in a greenhouse. The knowledge would be helpful in managing grasslands and restoring the degraded C. squarrosa grassland. Although there was neither inter- nor intra-specific competition between the two species when no nutrients were added, intra-specific competition of C. squarrosa was observed and increased with increased nutrient availability and more sulfur (S) was allocated to the aboveground partition of the plant. Relative competitive ability of C. squarrosa was greater than that of S. grandis when nutrients were supplied regardless of S. There was no significant difference between shoot and root competition based on dry matter yields. However, root competition was significantly greater than that of shoot based on S uptake under all treatments. A significant interaction was not observed between shoot and root competition. Therefore, nutrients addition benefits the restoration of degraded grassland of C. squarrosa, which may not exclude S. grandis. Also productivity and forage quality of the community will be increased. (C) 2007 Elsevier Ltd. All rights reserved.

An inventory of forest relicts in the pastures of Southern Tibet (Xizang AR, China)

An inventory of isolated tree stands surrounded by desert pastures in Southern Tibet (A.R. Xizang, China) revealed more than 50 sites with vigorous trees of Juniperus convallium Rehder & E.H. Wilson and Juniperus tibetica Kom and additional more than 10 records where juniper trees had been destroyed between 1959-1976. The tree stands are not restricted to any specific habitat, and occur within an area stretching 650 km westwards from the current forest border of Southern Tibet. The trees are religious landmarks of the Tibetan Buddhists. The highest trees were found at an elevation of 4,860 m. Vegetation records, rainfall correlations and temperature data collected by local climate stations and successful reforestation trials since 1999 indicate that forest relicts fragmented through human interference could regenerate if current cattle grazing and deforestation practices are halted. The drought line of Juniperus forests in Southern Tibet is approximately 200-250 mm/a. A first pollen diagram from Lhasa shows forest decline associated with the presence of humans since at least 4,600 yr BP. The currently degraded commons developed in the last 600 yr. To date, no findings of remains of ancient forests in the Central Tibetan Highlands of the Changtang have been reported.

Short-term effect of targeted placements of sheep excrement on grassland in inner mongolia on soil and plant parameters

Grazing animal excrement plays an important role in nutrient cycling and redistribution in grazing ecosystems, due to grazing in large areas and return in small areas. To elucidate the changes to the soil and pasture caused by sheep urine, fresh dung, and compost patches, a short- term field experiment using artificially placed pats was set up in the autumn of 2003 in the Inner Mongolian steppe. Urine application significantly increased soil pH during the first 32 days in soil layers at depths of both 0 - 5 cm and 5 - 15 cm. Rapid hydrolysis of urea gave large amounts of urine- nitrogen ( N) as ammonium ( NH4+) in soil extracts and was followed by apparent nitrification from day 2. Higher inorganic N content in the urine- treated soil was found throughout the experiment compared with the control. No significant effects of sheep excrement on soil microbial carbon ( C) and soil microbial N was found, but microbial activities significantly increased compared with the control after application of sheep excrement. Forty- six percent of dung- N and 27% of compost- N were transferred into vegetation after the experiment. The results from this study suggest that large amounts of nutrients have been lost from the returned excrement patches in the degraded grassland of Inner Mongolia, especially from sheep urine- N.

Population dynamics and responses to management of plateau pikas Ochotona curzoniae

1. Plateau pikas Ochotona curzoniae are considered a pest species on the Tibetan Plateau because they compete with livestock for forage and their burrowing could contribute to soil erosion. The effectiveness of pest control programmes in Tibet has not been measured, and it is not known whether changes in livestock management have exacerbated problems with plateau pikas or compromised their control. This study measured the impact of control programmes and livestock management for forage conservation on populations of plateau pikas in alpine meadow in Naqu District, central Tibet, during 2004 and 2005.2. Current techniques for controlling plateau pikas in spring cause large reductions in abundance, but high density-dependent rates of increase result in no differences between treated and untreated populations by the following autumn. Rates of increase from spring to autumn are not influenced by standing plant biomass or concurrent grazing by yaks Bos grunniens and Tibetan sheep Ovis aries.3. In autumn there was significantly lower biomass outside fenced areas with year-round livestock grazing compared with inside fenced areas with equivalent or higher numbers of plateau pikas but predominantly winter grazing by livestock. Inside fenced areas, control of plateau pikas in spring produced no detectable effect on standing plant biomass at the end of the following summer compared with uncontrolled populations of plateau pikas.4. Regardless of their initial density, populations of plateau pikas declined rapidly over winter outside fenced areas where there was very low standing plant biomass in autumn. However, inside fenced areas with higher plant biomass in autumn, low-density populations of plateau pikas declined more slowly than high-density populations.5. Synthesis and applications. Current control programmes have limited effect because populations of plateau pikas can recover in one breeding season. There was no apparent increase in forage production in areas where plateau pikas were controlled. However, plateau pikas appear to benefit from changes in grazing management, with low-density populations declining less over winter inside fenced areas than elsewhere. It was not evident that control programmes are warranted or that they will improve the livelihoods of Tibetan herders.

Responses of Artemisia frigida Willd. (Compositae) and Leymus chinensis (Trin.) Tzvel. (Poaceae) to sheep saliva

Although studies show that grazing and browsing by herbivores have marked effects on host plants, the mechanisms remain unclear. The objective of this study is to determine the effects of sheep saliva on host plant growth. Sheep saliva was manually applied to clipped plants of two different life forms, a semi-shrub, Artemisia frigida Willd., and a herbaceous species, Leymus chinensis (Trin.) Tzevel. The results showed that sheep saliva significantly enhanced aboveground net primary productivity (ANPP) and the ratio of ANPP to belowground net primary productivity (BNPP) for both species. This indicated that sheep saliva promotes aboveground compensatory growth and allocation of photosynthate to aboveground for both plant species. Sheep saliva stimulated only tillering of L. chinensis. Regardless of saliva application, clipping significantly decreased BNPP and plant height, but significantly increased the number of branches or tillers for both plant species. The relative growth rates (RGRs) on both species were significantly greater after clipping with saliva compared with control and clipping without saliva treatments. In addition, RGR of the herbaceous species L. chinensis was faster than that of the semi-shrub A. frigida after application of saliva. (c) 2006 Elsevier Ltd. All rights reserved.

Effects of forage composition and growing season on methane emission from sheep in the Inner Mongolia steppe of China

Understanding the effects of dietary composition on methane (CH4) production of sheep can help us to understand grassland degradation resulting in an increase of CH4 emission from ruminant livestock and its resulting significance affecting CH4 source/sink in the grazing ecosystem. The objective of this study was to investigate the effect of forage composition in the diet of sheep in July and August on CH4 production by sheep in the Inner Mongolia steppe. The four diet treatments were: (1) Leymus chinensis and Cleistogenes squarrosa (LC), (2) Leymus chinensis, Cleistogenes squarrosa and concentrate supplementation (LCC), (3) Artemisia frigida and Cleistogenes squarrosa (AC), and (4) Artemisia frigida, Cleistogenes squarrosa and concentrate supplementation (ACC). CH4 production was significantly lower in July than in August (31.4 and 36.2 g per sheep-unit per day, respectively). The daily average CH4 production per unit of digestive dry matter (DM), organic matter (OM) and neutral detergent fiber (NDF) increased by 10.9, 11.2 and 42.1% for the AC diet compared with the LC diet, respectively. Although concentrate supplementation in both the AC and LC diets increased total CH4 production per sheep per day, it improved sheep productivity and decreased CH4 production by 14.8, 12.5 and 14.8% per unit of DM, OM and NDF digested by the sheep, respectively. Our results suggested that in degraded grassland CH4 emission from sheep was increased and concentrate supplementation increased diet use efficiency. Sheep-grazing ecosystem seems to be a source of CH4 when the stocking rate is over 0.5 sheep-units ha(-1) during the growing season in the Inner Mongolia steppe.

Morphological and reproductive response of Caragana microphylla to different stocking rates

Understanding the effect on host plants of defending against herbivores is important in grazing ecology and grassland management. In this study, the morphological and reproductive responses of Caragana microphylla Lam. to grazing sheep were investigated using a 15-year grazing experiment with six stocking rates in the Inner Mongolia steppe of China. Plant height, rachis length, leaflet size, and number of pods decreased significantly, whereas spine density and length increased significantly with increased stocking rate. Significant negative correlations were observed between production of vegetative and reproductive organs and defensive organs, indicating that it is costly for C. microphylla to defend against herbivores and that morphological miniaturization and a tradeoff between production and defense were main responses of C. microphylla to herbivores. (c) 2006 Elsevier Ltd. All rights reserved.

Short-term effects of sheep excrement on carbon dioxide, nitrous oxide and methane fluxes in typical grassland of Inner Mongolia

Excrement patches of grazing animals play an important role in greenhouse gas (GHG) fluxes due to the high nitrogen (N) and available carbon (C) deposited in small areas, but little information is available for the effect of excrement in the Inner Mongolian grassland (43 26 degrees N, 116 degrees 40'E). To elucidate the effect of grazing sheep urine, fresh dung and compost on fluxes of methane (CH4), carbon dioxide (CO2), and nitrous oxide (N2O), a short-term field study (65 days) was carried out in the typical grassland of Inner Mongolia with the optimised closed chamber/GC technique. Compared with the control, cumulative net CH4 consumption decreased 36, 31, and 18% from urine, fresh dung, and compost plots, respectively; net CO2-C output increased by 6.5, 1.5, and 1.2% from urine, fresh dung, and compost treated soil, respectively; about three times as much N2O-N was emitted from urine and the fresh dung treatments during 65 days. Nitrous oxide emission was positively correlated with CO, emission (R = 0.691, P < 0.01) and water-filled pore space (R = 0.698, P < 0.01). The percentages of N2O-N loss of applied-N were 0.44 and 1.05% for urine and fresh dung, respectively. Our results suggest that in autumn in the degraded grassland of Inner Mongolia, the effect of sheep excrement may be ignored when evaluating the total GHG emissions.

The effect of supplementary feeds on the bodyweight of yaks in cold season

The present study was conducted to determine the effects of supplementary feeds, oat hay (OH), highland barley straw (HBS) and multi-nutrient blocks supplementation (UMMB) on reducing liveweight losses of both yak cows and calves grazed on low quality pastures during cold season. The trials of OH and HBS supplementation were conducted by using completely random design on 104 yak cows between 6 and 12 years of age as the following treatments: pure grazing (41 animals, body weight 230 67 kg) as control (CK); grazing+1.5 kg DM of OH per head daily (30 animals, body weight 216 28 kg); gazing. 1.5 kg DM of HBS per head daily (33 animals, body weight 221 34 kg). The trial of UMMB was conducted on three types of yaks, 1-year calves (8-12 months old, body weight 61.1 6.9 kg), 2-year calves (18-24 months old, 98.0 11.3 kg) and yak cows (164.5 27.1 (S.D.) kg) with 20 animals in control group (CK) and 20 animals in supplement group for each type by using completely random design as the following treatments: pure grazing for CK group; grazing+ 150, 250 and 500 g UMMB per day averagely for 1-year calf, 2-year calf and cow at night. The results indicate that the animals supplemented with oat hay received body weight gain (32 20.7 g day(-1)), while those supplemented with highland barley straw still suffered from body weight loss (-56.7 39.3 a day(-1)); UMMB supplementation can decrease the body weight loss by 109.7%, 86.6% and 63.4% for the 1-year calves, 2-year calves and yak cows, respectively, as compared with pure grazing. Around US$1.60 output can be achieved on the basis of US$1 input for UMMB supplementation in the farming systems of the 1-year calves, 2-year calves and yak cows, while US$1 input can produce US$1.55 and 1.14 output for OH and FIBS supplementations, respectively, in yak cows' farming system. It can be preliminary concluded that UMMB supplementation was the most economic way to alleviate body weight loss of grazing yaks over cold season, and the higher productive returns were obtained from OH supplementation for grazing yak cows during winter/spring months. © 2004 Elsevier B.V All rights reserved.

Local warming about 1.3 degrees C in alpine meadow has no effect on root vole (Microtus oeconomus L.) population during winter

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 m(2) 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 x 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 degrees 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 degrees C warming of soil and air introduced locally and on small (several m(2)) scale, in the alpine meadow habitat in winter period, has possibly no effect on root vole numbers and biomass.

Carbon dioxide exchange between the atmosphere and an alpine meadow ecosystem on the Qinghai-Tibetan Plateau, China

We used the eddy covariance method to measure the M exchange between the atmosphere and an alpine meadow ecosystem (37degrees29-45'N, 101degrees12-23'E, 3250m a.s.l.) on the Qinghai-Tibetan Plateau, China in the 2001 and 2002 growing seasons. The maximum rates Of CO2 uptake and release derived from the diurnal course Of CO2 flux (FCO2) were -10.8 and 4.4 mumol m(-2) s(-1), respectively, indicating a relatively high net carbon sequestration potential as compared to subalpine coniferous forest at similar elevation and latitude. The largest daily CO2 uptake was 3.9 g cm(-2) per day on 7 July 2002, which is less than half of those reported for lowland grassland and forest at similar latitudes. The daily CO2 uptake during the measurement period indicated that the alpine ecosystem might behave as a sink of atmospheric M during the growing season if the carbon lost due to grazing is not significant. The daytime CO2 uptake was linearly correlated with the daily photosynthetic photon flux density each month. The nighttime averaged F-CO2 showed a positive exponential correlation with the soil temperature, but apparently negative correlation with the soil water content. (C) 2004 Elsevier B.V. All rights reserved.