Elevational diversity of terrestrial rainforest herbs: when the whole is less than the sum of its parts

We studied the species richness of herbaceous terrestrial plant species along an elevational gradient at 250-2425m a.s.l. in evergreen tropical forest in Central Sulawesi, Indonesia. We recorded 302 species belonging to 51 families. Ferns and lycophytes contributed 62% of the species, followed by monocots with 24% and dicots with 14%. Overall herb species richness did not show any particular relation with elevation, while the richness of ferns increased significantly with elevation, monocots did not show a pattern, and dicots showed a hump-shaped pattern with maximum richness at 1800m. These patterns in turn were only partly reflected in the patterns of the individual plant families making up each group. The independence of different taxa was also reflected in their relationships to environmental factors (temperature, precipitation, and area): although, each single family was related to one or several factors, at the group level and at the overall level these trends were lost. These results show that interpreting diversity at higher taxonomic level may overlook important information at the family level and raises the biologically intriguing question whether overall patterns of diversity result from a random accumulation of group-specific patterns or if there is some interaction between groups (e.g., via competition and niche-pre-emption

The Role of Low Soil Temperature for Photosynthesis and Stomatal Conductance of Three Graminoids From Different Elevations

In high-elevation grasslands, plants can encounter periods with high air temperature while the soil remains cold, which may lead to a temporary mismatch in the physiological activity of leaves and roots. In a climate chamber experiment with graminoid species from three elevations (4400, 2400, and 250 m a.s.l.), we tested the hypothesis that soil temperature can influence photosynthesis and stomatal conductance independently of air temperature. Soil monoliths with swards of Kobresia pygmaea (high alpine), Nardus stricta (lower alpine), and Deschampsia flexuosa (upper lowland) were exposed to soil temperatures of 25, 15, 5, and -2°C and air temperatures of 20 and 10°C for examining the effect of independent soil and air temperature variation on photosynthesis, leaf dark respiration, and stomatal conductance and transpiration. Soil frost (-2°C) had a strong negative effect on gas exchange and stomatal conductance in all three species, independent of the elevation of origin. Leaf dark respiration was stimulated by soil frost in D. flexuosa, but not in K. pygmaea, which also had a lower temperature optimum of photosynthesis. Soil cooling from 15 to 5°C did not significantly reduce stomatal conductance and gas exchange in any of the species. We conclude that all three graminoids are able to maintain a relatively high root water uptake in cold, non-frozen soil, but the high-alpine K. pygmaea seems to be especially well adapted to warm shoot – cold root episodes, as it has a higher photosynthetic activity at 10 than 20°C air temperature and does not up-regulate leaf dark respiration upon soil freezing, as was observed in the grasses from warmer climates

Evapotranspiration and water balance of high-elevation grassland on the Tibetan Plateau

High-elevation grasslands of the Cyperaceae Kobresia pygmaea cover nearly half a million km2 on the Tibetan Plateau. As a consequence of climate change, precipitation patterns in this monsooninfluenced region may change with possible consequences for grassland productivity. Yet, not much is known about the water cycle in this second largest alpine ecosystem of the world. We measured the evapotranspiration of a high-elevation Kobresia pasture system at 4400 m a.s.l. in the south-eastern part of the plateau in two summers using three different approaches, weighable micro-lysimeters, eddy covariance measurements, and water balance modeling with the soil–plant–atmosphere transfer model SEWAB. In good agreement among the three approaches, we found ET rates of 4–6 mm d-1 in moist summer periods (June–August) and 2mmd-1 in dry periods, despite the high elevation and a leaf area index of only 1. Measured ET rates were comparable to rates reported from alpine grasslands at 1500–2500 m a.s.l. in temperate mountains, and also matched ET rates of managed lowland grasslands in the temperate zone. At the study site with 430 mm annual precipitation, low summer rainfall reduced ET significantly and infiltration into the subsoil occurred only in moist periods. Our results show that the evapotranspiration of high-elevation grasslands at 4400 m can be as high as in lowland grasslands despite large elevational changes in abiotic and biotic drivers of ET, and periodic water shortage is likely to influence large parts of the Tibetan Kobresia pastures

The Kobresia pygmaea ecosystem of the Tibetan highlands – Origin, functioning and degradation of the world's largest pastoral alpine ecosystem: Kobresia pastures of Tibet

With 450,000 km2 Kobresia (syn. Carex) pygmaea dominated pastures in the eastern Tibetan highlands are the world's largest pastoral alpine ecosystem forming a durable turf cover at 3000–6000 m a.s.l. Kobresia's resilience and competitiveness is based on dwarf habit, predominantly below-ground allocation of photo assimilates, mixture of seed production and clonal growth, and high genetic diversity. Kobresia growth is co-limited by livestock-mediated nutrient withdrawal and, in the drier parts of the plateau, low rainfall during the short and cold growing season. Overstocking has caused pasture degradation and soil deterioration over most parts of the Tibetan highlands and is the basis for this man-made ecosystem. Natural autocyclic processes of turf destruction and soil erosion are initiated through polygonal turf cover cracking, and accelerated by soil-dwelling endemic small mammals in the absence of predators. The major consequences of vegetation cover deterioration include the release of large amounts of C, earlier diurnal formation of clouds, and decreased surface temperatures. These effects decrease the recovery potential of Kobresia pastures and make them more vulnerable to anthropogenic pressure and climate change. Traditional migratory rangeland management was sustainable over millennia, and possibly still offers the best strategy to conserve and possibly increase C stocks in the Kobresia turf

Pasture degradation modifies the water and carbon cycles of the Tibetan highlands

The Tibetan Plateau has a significant role with regard to atmospheric circulation and the monsoon in particular. Changes between a closed plant cover and open bare soil are one of the striking effects of land use degradation observed with unsustainable range management or climate change, but experiments investigating changes of surface properties and processes together with atmospheric feedbacks are rare and have not been undertaken in the world's two largest alpine ecosystems, the alpine steppe and the Kobresia pygmaea pastures of the Tibetan Plateau. We connected measurements of micro-lysimeter, chamber, 13C labelling, and eddy covariance and combined the observations with land surface and atmospheric models, adapted to the highland conditions. This allowed us to analyse how three degradation stages affect the water and carbon cycle of pastures on the landscape scale within the core region of the Kobresia pygmaea ecosystem. The study revealed that increasing degradation of the Kobresia turf affects carbon allocation and strongly reduces the carbon uptake, compromising the function of Kobresia pastures as a carbon sink. Pasture degradation leads to a shift from transpiration to evaporation while a change in the sum of evapotranspiration over a longer period cannot be confirmed. The results show an earlier onset of convection and cloud generation, likely triggered by a shift in evapotranspiration timing when dominated by evaporation. Consequently, precipitation starts earlier and clouds decrease the incoming solar radiation. In summary, the changes in surface properties by pasture degradation found on the highland have a significant influence on larger scales

The Kobresia pygmaea ecosystem of the Tibetan highlands – Origin, functioning and degradation of the world's largest pastoral alpine ecosystem

Kobresia pastures in the eastern Tibetan highlands occupy 450000 km2 and form the world’s largest pastoral alpine ecosystem. The main constituent is an endemic dwarf sedge, Kobresia pygmaea, which forms a lawn with a durable turf cover anchored by a felty root mat, and occurs from 3000 m to nearly 6000 m a.s.l. The existence and functioning of this unique ecosystem and its turf cover have not yet been explained against a backdrop of natural and anthropogenic factors, and thus its origin, drivers, vulnerability or resilience remain largely unknown. Here we present a review on ecosystem diversity, reproduction and ecology of the key species, pasture health, cycles of carbon (C), water and nutrients, and on the paleo-environment. The methods employed include molecular analysis, grazing exclusion, measurements with micro-lysimeters and gas exchange chambers, 13C and 15N labelling, eddy-covariance flux measurements, remote sensing and atmospheric modelling. The following combination of traits makes Kobresia pygmaea resilient and highly competitive: dwarf habit, predominantly below-ground allocation of photo assimilates, mixed reproduction strategy with both seed production and clonal growth, and high genetic diversity. Growth of Kobresia pastures is co-limited by low rainfall during the short growing season and livestock-mediated nutrient withdrawal. Overstocking has caused pasture degradation and soil deterioration, yet the extent remains debated. In addition, we newly describe natural autocyclic processes of turf erosion initiated through polygonal cracking of the turf cover, and accelerated by soil-dwelling endemic small mammals. The major consequences of the deterioration of the vegetation cover and its turf include: (1) the release of large amounts of C and nutrients and (2) earlier diurnal formation of clouds resulting in (3) decreased surface temperatures with (4) likely consequences for atmospheric circulation on large regional and, possibly global, scales. Paleo-environmental reconstruction, in conjunction with grazing experiments, suggests that the present grazing lawns of Kobresia pygmaea are synanthropic and may have existed since the onset of pastoralism. The traditional migratory rangeland management was sustainable over millennia and possibly still offers the best strategy to conserve, and possibly increase, the C stocks in the Kobresia turf, as well as its importance for climate regulation