Abstract
We explored patterns of plant species richness and composition along an elevational gradient (4,985–5,685 m a.s.l.) on Buddha Mountain, 100 km northwest of Lhasa, Tibet. We recorded the presence of plants and lichens in 1-m2 quadrats separated by 25-m elevational intervals (174 quadrats in 29 elevational bands) along a vertical transect with a SE aspect. We recorded 143 total species, including 107 angiosperms, 2 gymnosperms, 27 lichens, and 7 mosses. We measured stone cover in each quadrat, and soil pH, C, N and C/N ratio from two randomly located samples collected from 10-cm depth within each band. C, N and C/N decreased with elevation, stoniness increased and soil pH did not change with altitude. We employed detrended correspondence analysis (DCA), canonical correspondence analysis (CCA) and generalized linear models (GLMs) to assess the relationships of species richness and species composition to the environment. The first two axes of the CCA biplot explained 87.7% of total variation in the species-environment relationship, and 27.7% of total variance of species data. The first CCA axis is associated with elevation, while the second axis is related to soil pH and stone cover. We also compared patterns in species richness against expectations from species pools interpolated from the literature. Total species richness was relatively constant between 4,985 and 5,400 m a.s.l. and declined continuously above 5,400 m a.s.l. Similar declining patterns were observed for forbs and graminoids. Cushion plants and lichens abundance exhibited a unimodal relationship with altitude while shrubs declined monotonically. Except for lichens, models derived from our observations and the literature were quite similar in shape. The proportion of the species pool represented in each elevational band increased as a function of elevation for non-vascular plants, but decreased markedly for vascular plants. Thus, vascular plants are more likely to be constrained by dispersal at higher elevations, resulting in more local endemism, while the relatively easily-dispersed high-elevation cryptogams have little local differentiation. Our comparative approach demonstrates that complex scale-dependent differences between life forms may underlie the apparent simplicity of elevational gradients. Furthermore, elevational gradients summarized from distributional notes cannot be assumed to be proxies for elevational gradients on individual mountain slopes.
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References
Baniya CB (2010) Vascular and cryptogam richness in the world’s highest alpine zone, Tibet. Mount Res Developm 30:275–281
Baniya CB, Solhøy T, Gauslaa Y, Palmer MW (2010) The elevation gradient of lichen species richness in Nepal. The Lichenologist 42:83–96
Birks HJB, Birks HH, Everson J, Jans H, Thorne D, Thorne M (2007) The AGS in Tibet 2005. Alpine Gardener 75:289–349
Brown JH (1981) Two decades of homage to Santa Rosalia: toward a general theory of diversity. Amer Zool 21:877–888
Bruun HH, Moen J, Virtanen R, Grytnes JA, Oksanen L, Angerbjörn A (2006) Effects of altitude and topography on species richness of vascular plants, bryophytes and lichens in alpine communities. J Veg Sci 17:37–46
Callaway RMR (2002) Positive interactions among alpine plants increase with stress. Nature 417:844
Chang DHS (1981) The vegetation zonation of the Tibetan Plateau. Mount Res Developm 1:29–48
Colwell RK, Lees DC (2000) The Mid-Domain Effect: Geometric constraints on the geography of species richness. Trends Ecol Evol 15:70–76
Cornell HV (1993) Unsaturated patterns in species assemblages: the role of regional processes in setting local species richness. In Ricklefs RE, Schluter D (eds) Species diversity in ecological communities: historical and geographical perspectives. Chicago University Press, Chicago, pp 243–252
de Vera JP, Horneck G, Rettberg P, Ott S (2004) The potential of the lichen symbiosis to cope with the extreme conditions of outer space II: germination capacity of lichen ascospores in response to simulated space conditions. Advances Space Res 33:1236–1243
Du Z (1992) A study of the altitudinal belt of vegetation in the southeastern part of the Qinghai-Xizang (Tibetan) Plateau. Braun-Blanquetia 8:84–86
Dvorský M, Doležal J, De Bello F, Klimešová J, Klimeš L (2011) Vegetation types of East Ladakh: species and growth form composition along main environmental gradients. Appl Veg Sci 14:132–147
Feuerer T, Hawksworth DL (2007) Biodiversity of lichens, including a world-wide analysis of checklist data based on Takhtajan’s floristic regions. Biodivers & Conservation 16:85–98
Grabherr G, Gottfried M, Gruber A, Pauli H (1995) Patterns and current changes in alpine plant diversity. In Chapin III FS, Körner C (eds) Arctic and alpine biodiversity: patterns, causes and ecosystem consequences. Ecological studies 113, Springer-Verlag, Berlin, pp 167–181
Grytnes JA, Vetaas OR (2002) Species richness and altitude: a comparison between null models and interpolated plant species richness along the Himalayan altitudinal gradient, Nepal. Amer Naturalist 159:294–304
Grytnes JA, Heegaard E, Ihlen PG (2006) Species richness of vascular plants, bryophytes, and lichens along an altitudinal gradient in western Norway. Acta Oecol 29:241–246
Harris N (2006) The elevation history of the Tibetan Plateau and its implications for the Asian monsoon. Palaeogeogr, Palaeoclimatol, Palaeoecol 241:4–15
Hertel H (1977) Gesteinsbewohnende Arten der Sammelgattung Lecidea (Lichenes) aus Zentral-, Ost- und Südasien. Eine erste Übersicht. Khumbu Himal 6:145–378
Herzschuh U, Birks HJB, Mischke S, Zhang C, Böhner JR (2010) A modern pollen–climate calibration set based on lake sediments from the Tibetan Plateau and its application to a late quaternary pollen record from the Qilian Mountains. J Biogeogr 37:752–766
Huston MA (1994) Biological diversity: the coexistence of species on changing landscapes. Cambridge University Press, Cambridge
Jin-Ting W (1992) A preliminary study on alpine vegetation in Qinghai-Xizang (Tibet) Plateau. Braun-Blanquetia 8:82–83
Jongman RHG, ter Braak JFT, Van Tongeren OFR (1995) Data analysis in community and landscape ecology. Cambridge University Press, Cambridge
Jürgen K, Kessler M, Robert RD (2006) What drives elevational patterns of diversity? A test of geometric constraints, climate and species pool effects for pteridophytes on an elevational gradient in Costa Rica. Global Ecol Biogeogr 15:358–371
Kessler M (2000) Altitudinal zonation of Andean cryptogam communities. J Biogeogr 27:275–282
Klein JA, Harte J, Zhao X-Q (2007) Experimental warming, not grazing, decreases rangeland quality on the Tibetan Plateau. Ecol Applications 17:541–557
Klimeš L, Doležal J (2010) An experimental assessment of the upper elevational limit of flowering plants in the western Himalayas. Ecography 33:590–596
Körner C (2000) Why are there global gradients in species richness? Mountains might hold the answer. Trends Ecol Evol 15:513–514
Körner C (2003) Alpine plant life: functional plant ecology of high mountain ecosystems. Springer Verlag, Berlin
Lepš J, Šmilauer P (2003) Multivariate analysis of ecological data using CANOCO. University of Cambridge, Cambridge
Li XJ (1985) Bryoflora of Xizang. Science Press, Beijing (In Chinese)
McCullagh P, Nelder JA (1989) Generalized linear models. Chapman and Hall, London
Miehe G (1988) Geoecological reconnaissance in the alpine belt of southern Tibet. GeoJournal 17:635–648
Miehe G (1991) Der Himalaya, eine multizonale Gebirgsregion. In Walter H, Breckle S (eds) Ökologie der gemäβigten und arktischen Zonen auβerhalb Euro-Nordasiens. Gustav Fischer, Stuttgart, pp 181–230
Miehe G, Miehe S, Schlütz F, Kaiser K, Duo L (2006) Palaeoecological and experimental evidence of former forests and woodlands in the treeless desert pastures of Southern Tibet (Lhasa, A. R. Xizang, China). Palaeogeogr Palaeoclimatol Palaeoecol 242:54–67
Ni J (2000) A simulation of biomes on the Tibetan Plateau and their responses to global climate change. Mount Res Developm 20:80–89
Nogués-Bravo D, Araujo MB, Romdal T, Rahbek C (2008) Scale effects and human impact on the elevational species richness gradients. Nature 453:216–219
O’Brien EM (1993) Climatic gradients in woody plant species richness: towards an explanation based on an analysis of Southern Africa’s woody flora. J Biogeogr 20:181–198
Odland A, Birks HJB (1999) The altitudinal gradient of vascular plant species richness in Aurland, western Norway. Ecography 22:548–566
Palmer MW (2006) Scale dependence of native and alien species richness in North American floras. Preslia 78:427–436
Palmer MW, Dixon PM (1990) Small-scale environmental heterogeneity and the analysis of species distributions along gradients. J Veg Sci 1:57–65
Pauli H, Gottfried M, Reiter K, Klettner C, Grabherr G (2007) Signals of range expansions and contractions of vascular plants in the high Alps: observations (1994–2004) at the GLORIA master site Schrankogel, Tyrol, Austria. Global Change Biol 13:147–156
Pavon NP, Hernandez-Trejo H, Rico-Gray V (2000) Distribution of plant life forms along an altitudinal gradient in the semi-arid valley of Zapotitlan, Mexico. J Veg Sci 11:39–42
Polunin O, Stainton JDA (2001) Concise flowers of the Himalaya. Oxford University Press, Oxford
R Development Core Team (2008) R: A language and environment for statistical computing version 2.8.1. R Foundation for Statistical Computing, Vienna
Rahbek C (1995) The elevational gradient of species richness: a uniform pattern? Ecography 18:200–205
Rahbek C (2005) The role of spatial scale and the perception of large-scale species-richness patterns. Ecol Lett 8:224–239
Sancho LG, de la Torre R, Horneck G, Ascaso C, de los Rios A, Pintado A, Wierzchos J, Schuster M (2007) Lichens survive in space: results from the 2005 LICHENS Experiment. Astrobiology 7:443–454
Shu J (1992) On the vegetation zonation of Qinghai-Xizang Plateau. Braun-Blanquetia 8:80–81
Spicer RA, Harris NBW, Widdowson M, Herman AB, Guo S, Valdes PJ, Wolfe JA, Kelley SP (2003) Constant elevation of southern Tibet over the past 15 million years. Nature 421:622–624
Stainton JDA (2001) Flowers of the Himalaya: a supplement. Oxford University Press, Oxford
Sumner ME (1999) Handbook of soil science. CRC Press, Boca Raton
ter Braak CJF (2002) CANOCO — a FORTRAN program for canonical community ordination by (partial) (detrended) (canonical) correspondence analysis, principal component analysis and redundancy analysis, Version 4.5. Biometris-Plant Research International, Wageningen
ter Braak CJF, Šmilauer P (2002) CANOCO reference manual and CanoDraw for Windows user’s guide: software for canonical community ordination (version 4.5). Microcomputer Power, Ithaca, NY
Theurillat JP, Schlüssel A, Geissler P, Guisan A, Velluti C, Wiget L (2003) Vascular plant and bryophyte diversity along elevational gradients in the Alps. In Nagy L, Grabherr G, Körner C, Thompson DBA (eds) Alpine biodiversity in Europe. Ecological Studies 167, Springer Verlag, Berlin, pp 185–193
Virtanen R, Crawley MJ (2010). Contrasting patterns in bryophyte and vascular plant species richness in relation to elevation, biomass and Soay sheep on St Kilda, Scotland. Pl Ecol Divers 3:77–85
Virtanen R, Dirnböck T, Dullinger S, Grabherr G, Pauli H, Staudinger M, Villar L (2003) Patterns in the plant species richness of European high mountain vegetation. In Nagy L, Grabherr G, Körner C, Thompson DBA (eds) Alpine biodiversity in Europe. Ecological Studies 167, Springer Verlag, Berlin, pp 149–172
Wang QJ, Liu JQ, Zhao XQ (2002) Patterns of plant species diversity in the Northeastern Tibetan Plateau, Qinghai, China. In Körner C, Spehn E (eds) Mountain biodiversity — a global assessment. Parthenon, New York, pp 149–153
Wang WY, Wang QJ, Li SX, Wang G (2006) Distribution and species diversity of plant communities along transect on the Northeastern Tibetan plateau. Biodivers & Conservation 15:1811–1828
Wang Z, Tang Z, Fang J (2007) Altitudinal patterns of seed plant richness in the Gaoligong Mountains, south-east Tibet, China. Diversity Distrib 13:845–854
Wei J-C, Jiang Y-M (1986) Lichens of Xizang. Science Press, Beijing (In Chinese)
Weilie C, Jinting W, Bosheng L (1992) The main types of vegetation and their distribution in Tibet. Braun-Blanquetia 8:77–79
Whittaker RJ, Willis KJ, Field R (2001) Scale and species richness: towards a general, hierarchical theory of species diversity. J Biogeogr 28:453–470
Wright DH (1983) Species-energy theory: an extension of species-area theory. Oikos 41:496–506
Wu C-7Y (1983–1987) Flora Xizangica. Vol. I–V. Science Press, Beijing (In Chinese)
Acknowledgements
We thank the Norwegian State Education Loan Fund (Lånekassen) for funding, the Norway-Tibet Network for partial travel support, and the Central Department of Botany, Tribhuvan University, Kathmandu, Nepal for leave to Baniya, CB. We also thank John Birks for his help in plant identifications and valuable suggestions after going through some earlier versions of this manuscript. Tsering, Cai Dong, Pubu, La Qiong, LaDuo, Droba, and Frode Falkenberg helped to complete this study. Thanks are also due to Kristian Hassel for helping with moss determinations, Per Magnus Jørgensen for helping identify lichens, Petr Šmilauer and three anonymous referees for their useful comments and suggestions. Thanks to Joachim Schmidt for the map of the study area. Michael Denslow provided linguistic editing.
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Baniya, C.B., Solhøy, T., Gauslaa, Y. et al. Richness and Composition of Vascular Plants and Cryptogams along a High Elevational Gradient on Buddha Mountain, Central Tibet. Folia Geobot 47, 135–151 (2012). https://doi.org/10.1007/s12224-011-9113-x
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DOI: https://doi.org/10.1007/s12224-011-9113-x