Causes for the unimodal pattern of biomass and productivity in alpine grasslands along a large altitudinal gradient in semi-arid regions

Questions How can we understand the limitations to plant growth at high altitudes? Our aim was to test the hypotheses that for alpine grasslands along a large altitudinal gradient in semi-arid regions, plant growth is mainly limited by drought at low altitudes but by low temperature at high altitudes, resulting in a unimodal pattern of biomass and productivity associated with an optimal combination of temperature and precipitation. Such knowledge is important to understanding the response of alpine ecosystems to climate change. Location We conducted a 5-yr livestock exclosure experiment along the south-facing slope of the Nyaiqentanglha Mountains, central Tibetan Plateau. Methods We measured above- and below-ground biomass, species richness, leaf δ13C and water potential, and related climate and soil variables across 42 fenced and unfenced quadrats near seven HOBO weather stations along the slope. The vegetation changed from alpine steppe-meadow at 4390–4500 m to alpine meadow at 4600–5210 m. Results Total above- and below-ground biomass across fenced and unfenced quadrats increased with increasing altitude up to 4950–5100 m, and then decreased above 5100 m. Altitudinal trends in leaf δ13C and water potential of dominant species also showed a unimodal pattern corresponding to that of vegetation biomass. Total above- and below-ground biomass as well as sedge above-ground biomass all showed a quadratic relationship with mean temperatures and the ratio of growing season precipitation (GSP) to ≥5 °C accumulated temperature (AccT; R2 = 0.83−0.88, P < 0.001). In general, above- and below-ground biomass increased with increasing water availability when the GSP/AccT ratio was lower than the threshold level of 0.80–0.84, but decreased when the GSP/AccT ratio was higher than this threshold level. No significant relationship was found between residuals of above-ground biomass and species richness after removing the effects of climate factors on both stand variables. Conclusions The results support our hypotheses, further suggesting a threshold of water limitation that is consistent with the model prediction over the Tibetan Plateau. Species richness per se appears to weakly affect community-level productivity. The response of alpine grasslands to climate warming may vary with altitude because of altitudinal shifts in factors limiting plant growth.

[1]  Robert H. Whittaker,et al.  VEGETATION OF THE SANTA CATALINA MOUNTAINS, ARIZONA. V. BIOMASS, PRODUCTION, AND DIVERSITY ALONG THE ELEVATION GRADIENT' , 1975 .

[2]  H. Lieth Modeling the Primary Productivity of the World , 1975 .

[3]  Graham D. Farquhar,et al.  Carbon isotope discrimination is positively correlated with grain yield and dry matter production in field-grown wheat , 1987 .

[4]  J. Eischeid,et al.  Precipitation Fluctuations over Northern Hemisphere Land Areas Since the Mid-19th Century , 1987, Science.

[5]  O. Sala,et al.  A Generalized Model of the Effects of Grazing by Large Herbivores on Grassland Community Structure , 1988, The American Naturalist.

[6]  W. Parton,et al.  Primary Production of the Central Grassland Region of the United States , 1988 .

[7]  J. Ellis,et al.  Stability of African pastoral ecosystems: alternate paradigms and implications for development , 1988 .

[8]  J. Ehleringer,et al.  Carbon Isotope Discrimination and Photosynthesis , 1989 .

[9]  F. Woodward,et al.  Evolutionary and Ecophysiological Responses of Mountain Plants to the Growing Season Environment , 1990 .

[10]  F. Woodward,et al.  Experimental Investigations on the Environmental Determination of δ13C at Different Altitudes , 1990 .

[11]  Robert W. Howarth,et al.  Nitrogen limitation on land and in the sea: How can it occur? , 1991 .

[12]  Peter D. Erskine,et al.  13C Natural Abundance in Plant Communities Along a Rainfall Gradient: a Biological Integrator of Water Availability , 1995 .

[13]  F. Woodward,et al.  Experiments on the causes of altitudinal differences in the leaf nutrient contents, size and δ13C of Alchemilla alpina , 1996 .

[14]  D. Tilman,et al.  Productivity and sustainability influenced by biodiversity in grassland ecosystems , 1996, Nature.

[15]  I. Burke,et al.  PRODUCTIVITY PATTERNS OF C3 AND C4 FUNCTIONAL TYPES IN THE U.S. GREAT PLAINS , 1997 .

[16]  Michael A. Huston,et al.  Hidden treatments in ecological experiments: re-evaluating the ecosystem function of biodiversity , 1997, Oecologia.

[17]  J. P. Grime,et al.  Benefits of plant diversity to ecosystems: immediate, filter and founder effects , 1998 .

[18]  F. Chapin,et al.  Consequences of changing biodiversity , 2000, Nature.

[19]  J. P. Grime,et al.  No consistent effect of plant diversity on productivity. , 2000, Science.

[20]  J. Frangi,et al.  Grassland biomass dynamics along an altitudinal gradient in the Pampa. , 2000 .

[21]  Christian Hitz,et al.  Below-ground and above-ground production of vegetational organic matter along a climosequence in alpine grasslands , 2001 .

[22]  Katherine L. Gross,et al.  WHAT IS THE OBSERVED RELATIONSHIP BETWEEN SPECIES RICHNESS AND PRODUCTIVITY , 2001 .

[23]  A. Knapp,et al.  Variation among biomes in temporal dynamics of aboveground primary production. , 2001, Science.

[24]  S. Leavitt,et al.  Leaf δ13C variability with elevation, slope aspect, and precipitation in the southwest United States , 2002, Oecologia.

[25]  Huazhong Zhu,et al.  ESTIMATED BIOMASS AND PRODUCTIVITY OF NATURAL VEGETATION ON THE TIBETAN PLATEAU , 2002 .

[26]  C. Körner,et al.  Carbon isotope discrimination by plants follows latitudinal and altitudinal trends , 1991, Oecologia.

[27]  C. Körner,et al.  A global survey of carbon isotope discrimination in plants from high altitude , 2004, Oecologia.

[28]  T. Boutton,et al.  Distribution of biomass of species differing in photosynthetic pathway along an altitudinal transect in southeastern wyoming grassland , 2004, Oecologia.

[29]  C. Körner The nutritional status of plants from high altitudes , 1989, Oecologia.

[30]  C. Field,et al.  Variation in foliar δ13C in Hawaiian Metrosideros polymorpha: a case of internal resistance? , 1990, Oecologia.

[31]  M. Oesterheld,et al.  Effect of grazing on community structure and productivity of a Uruguayan grassland , 2005, Plant Ecology.

[32]  Yude Pan,et al.  Leaf area index and net primary productivity along subtropical to alpine gradients in the Tibetan Plateau , 2004 .

[33]  S. Macko,et al.  Natural abundance of 13C and 15N in C3 and C4 vegetation of southern Africa: patterns and implications , 2004 .

[34]  Sandra A. Brown,et al.  Root biomass along subtropical to alpine gradients: global implication from Tibetan transect studies , 2005 .

[35]  C. Lortie,et al.  LINKING PATTERNS AND PROCESSES IN ALPINE PLANT COMMUNITIES: A GLOBAL STUDY , 2005 .

[36]  P. Reich,et al.  Nitrogen limitation constrains sustainability of ecosystem response to CO2 , 2006, Nature.

[37]  K. Yoda,et al.  Climate and vegetation in China II. Distribution of main vegetation types and thermal climate , 1989, Ecological Research.

[38]  K. Nadrowski,et al.  Variation of precipitation and its effect on phytomass production and consumption by livestock and large wild herbivores along an altitudinal gradient during a drought, South Gobi, Mongolia , 2006 .

[39]  G. Wieser,et al.  Trees at their Upper Limit , 2007 .

[40]  A. Troumbis,et al.  Biodiversity and ecosystem functioning: reconciling the results of experimental and observational studies , 2007 .

[41]  James B Grace,et al.  Does species diversity limit productivity in natural grassland communities? , 2007, Ecology letters.

[42]  Qi Wang,et al.  Effects of altitude on plant-species diversity and productivity in an alpine meadow, Qinghai-Tibetan plateau , 2007 .

[43]  C. Peng,et al.  Leaf δ13C reflects ecosystem patterns and responses of alpine plants to the environments on the Tibetan Plateau , 2008 .

[44]  J. Larsen,et al.  Towards a physical description of habitat: quantifying environmental adversity (abiotic stress) in temperate forest and woodland ecosystems , 2009 .

[45]  Jingyun Fang,et al.  Aboveground biomass in Tibetan grasslands , 2009 .

[46]  C. Daly,et al.  Correlations between net primary productivity and foliar carbon isotope ratio across a Tibetan ecosystem transect , 2009 .

[47]  C. Daly,et al.  Leaf life span as a simple predictor of evergreen forest zonation in China , 2009 .

[48]  Lin Jiang,et al.  Species diversity and productivity: why do results of diversity‐manipulation experiments differ from natural patterns? , 2009 .

[49]  Yanhong Tang,et al.  Altitudinal variation of ecosystem CO2 fluxes in an alpine grassland from 3600 to 4200 m , 2009 .

[50]  Anne Zemmrich,et al.  Driving environmental factors and the role of grazing in grassland communities: A comparative study along an altitudinal gradient in Western Mongolia , 2010 .

[51]  Jingyun Fang,et al.  Environmental factors covary with plant diversity-productivity relationships among Chinese grassland sites , 2010 .

[52]  Sassan Saatchi,et al.  Introduction: Elevation gradients in the tropics: laboratories for ecosystem ecology and global change research , 2010 .

[53]  H. Prins,et al.  Biomass and diversity of dry alpine plant communities along altitudinal gradients in the Himalayas , 2011, Journal of Plant Research.