Orchid species richness along Himalayan elevational gradients
Kamal P. Acharya
Department of Biology, University of Bergen, PO Box 7803, N-5020, Bergen, Norway
Present address: Department of Biology, Norwegian University of Science and Technology, 7491, Trondheim, Norway
Search for more papers by this authorCorresponding Author
Ole R. Vetaas
Department of Geography, University of Bergen, Post Box 7802, N-5020 Bergen, Norway
Ole R. Vetaas, Department of Geography, University of Bergen, PO Box 7802, N-5020 Bergen, Norway.E-mail: [email protected]Search for more papers by this authorH. J. B. Birks
Department of Biology, University of Bergen, PO Box 7803, N-5020, Bergen, Norway
School of Geography and the Environment, University of Oxford, Oxford OX1 3QY, UK
Environmental Change Research Centre, University College London, London WC1E 6BT, UK
Search for more papers by this authorKamal P. Acharya
Department of Biology, University of Bergen, PO Box 7803, N-5020, Bergen, Norway
Present address: Department of Biology, Norwegian University of Science and Technology, 7491, Trondheim, Norway
Search for more papers by this authorCorresponding Author
Ole R. Vetaas
Department of Geography, University of Bergen, Post Box 7802, N-5020 Bergen, Norway
Ole R. Vetaas, Department of Geography, University of Bergen, PO Box 7802, N-5020 Bergen, Norway.E-mail: [email protected]Search for more papers by this authorH. J. B. Birks
Department of Biology, University of Bergen, PO Box 7803, N-5020, Bergen, Norway
School of Geography and the Environment, University of Oxford, Oxford OX1 3QY, UK
Environmental Change Research Centre, University College London, London WC1E 6BT, UK
Search for more papers by this authorAbstract
Aim This study explores the relationship between orchid species richness along elevational gradients in the central and eastern Himalaya. Specifically, we aimed to test predictions of maximum richness at different elevational levels for orchid species geographically restricted to central and eastern Himalaya (endemics), and total orchid richness, including different life forms. We then compared and assessed climatic factors together with area as causal explanations for the observed patterns.
Location Nepal (100–5200 m a.s.l.), Bhutan (100–5200 m a.s.l.), and adjacent regions of India, i.e. Sikkim and Darjeeling.
Methods We used published data on the distribution of orchids to interpolate presence between maximum and minimum recorded elevations in 100-m elevational zones. These patterns were compared with previously published patterns for other plant groups, which were derived in the same way. Descriptive regressions with metres above sea level (m a.s.l.) as the independent variable were performed using nonparametric generalized additive models (GAMs), whereas tests of the diversity models were carried out using parametric generalized linear models (GLMs).
Results A hump-shaped relationship between orchid species richness and elevation was observed in Nepal and Bhutan, with maximum richness at 1600 m a.s.l. corresponding to c. 16 °C mean annual temperature. The richness of orchid species that are endemic to central and eastern Himalaya showed a bimodal pattern. Richness of endemic epiphytes peaked at elevation zones similar to those for total richness, but the peak in numbers of endemic terrestrial species occurred at a higher elevation.
Main conclusions This study shows that climatic variables (temperature and precipitation) provide a good potential explanation for orchid species richness along the Himalayan elevational gradient, even when the effect of area is considered. The principal elevational peak in richness of central and eastern Himalayan endemics was consistent with the peak in total orchid richness, but maximum terrestrial endemic richness occurred at higher elevations.
References
- Acharya, K.P. & Rokaya, M.B. (2005) Ethnobotanical survey of medicinal plants traded in the streets of Kathmandu valley. Scientific World, 3, 44–48.
- Ackerman, J.D., Trejo-Torres, J.C. & Crespo-Chury, Y. (2007) Orchids of the West Indies: predictability of diversity and endemism. Journal of Biogeography, 34, 779–786.
- Anon. (1997) Climatological records of Nepal, 1976–1997. HMG Department of Hydrology and Meteorology, Kathmandu, Nepal.
- Anon. (2000) Land use and statistics section: a brief introduction to water resources, water use and water resource management in Bhutan. Ministry of Agriculture, Thimpu, Bhutan.
- Baillie, I.C. & Norbu, C. (2004) Climate and other factors in the development of river and interfluvial profiles in Bhutan, Eastern Himalayas. Journal of Asian Earth Science, 22, 539–553.
- Banerji, M.L. (1963) Outline of Nepal phytogeography. Vegetatio, 11, 288–296.
- Banerji, M.L. (1978) Orchids of Nepal, reprinted 1982. Bishen singh Mahendra pal Singh, Dehradun, India.
- Baniya, C.B., Solhøy, T., Gauslaa, Y. & Palmer, M.N. (2010) The elevation gradient of lichen species richness in Nepal. The Lichenologist, 42, 83–96.
- Beug, H.J. & Miehe, G. (1999) Vegetation history and human impact in the eastern central Himalaya (Langtang and Helambu, Nepal). Dissertationes Botanicae, 318, 1–98.
- Bhattarai, K.R. & Vetaas, O.R. (2003) Variation in plant species richness of different life forms along a subtropical elevation gradient in the Himalayas, east Nepal. Global Ecology and Biogeography, 12, 327–340.
- Bhattarai, K.R., Vetaas, O.R. & Grytnes, J.A. (2004) Fern species richness along a central Himalayan elevational gradient, Nepal. Journal of Biogeography, 31, 389–400.
- Blanchard, M.G. & Runkle, E.S. (2006) Temperature during the day, but not during the night, controls flowering of Phalaenopsis orchids. Journal of Experimental Botany, 57, 4043–4049.
- Cardelus, C.L., Colwell, R.K. & Watkins, J.E. (2006) Vascular epiphyte distribution patterns: explaining the mid-elevation richness peak. Journal of Ecology, 94, 144–156.
- Carpenter, C. (2005) The environmental control of plant species density on a Himalayan elevation gradient. Journal of Biogeography, 32, 999–1018.
- Chaudhary, R.P. (1998) Biodiversity of Nepal: status and conservation. S. Devi, India.
-
Crawley, M.J. (2005) Statistics: an introduction using R. John Wiley & Sons, Chichester, UK.
10.1002/9781119941750 Google Scholar
- Dobremez, J.F. (1976) Le Népal, écologie et biogéographie. Centre National de la Pecherche Scientifique, Paris.
- Dressler, R.L. (1993) Phylogeny and classification of the orchid family. Dioscorides Press, Portland, OR.
- Fossa, A.M. (2004) Biodiversity patterns of vascular plant species in mountain vegetation in the Faroe Islands. Diversity and Distributions, 10, 217–223.
- Grau, O., Grytnes, J.A. & Birks, H.J.B. (2007) A comparison of altitudinal species richness patterns of bryophytes with other plant groups in Nepal, Central Himalaya. Journal of Biogeography, 34, 1907–1915.
-
Grytnes, J.A. &
McCain, C.M. (2007) Elevational trends in biodiversity.
Encyclopedia of Biodiversity, Vol. II (ed. by
S. Levin), pp. 1–8. Elsevier, Oxford.
10.1016/B978-012226865-6/00503-1 Google Scholar
- Grytnes, J.A. & Vetaas, O.R. (2002) Species richness and altitude: a comparison between null models and interpolated plant species richness along the Himalayan altitudinal gradient, Nepal. The American Naturalist, 159, 294–304.
- Grytnes, J.A., Heegard, E. & Ihlen, P.G. (2006) Species richness of vascular plants, bryophytes, and lichens along an altitudinal gradient in western Norway. Acta Oecologica, 29, 241–246.
- Hara, H., Strearn, W.T. & Williams, L.H.J. (1978) An enumeration of flowering plants of Nepal. British Museum (Natural History), London.
- Hastie, T.J. & Tibshirani, R.J. (1990) Generalized additive models. Chapman & Hall, London.
- Hawkins, B.A., Field, R., Cornell, H.V., Currie, D.J., Guégan, J.F., Kaufman, D.M., Kerr, J.T., Mittelbach, G.G., Oberdorff, T., O’Brien, E.M., Porter, E.E. & Turner, J.R.G. (2003) Energy, water and broad-scale geographic patterns of species richness. Ecology, 84, 3105–3117.
- Hunter, M.L. & Yonzon, P. (1993) Altitudinal distributions of birds, mammals, people, forests, and parks in Nepal. Conservation Biology, 7, 420–423.
- Jacquemyn, H., Micheneau, C., Roberts, D.L. & Pailler, T. (2005) Elevational gradients of species diversity, breeding system and floral traits of orchid species on Réunion Island. Journal of Biogeography, 32, 1751–1761.
- Karger, D.N., Kluge, J., Krömer, T., Hemp, A., Lehnert, M. & Kessler, M. (2011) The effect of area on local and regional elevational patterns of species richness. Journal of Biogeography, 38, 1177–1185.
- Klanderud, K. & Birks, H.J.B. (2003) Recent increases in species richness and shifts in altitudinal distributions of Norwegian mountain plants. The Holocene, 13, 1–6.
- Kluge, J., Kessler, M. & Dunn, R.R. (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 Ecology and Biogeography, 15, 358–371.
- Körner, C. (2000) Why are there global gradients in species richness? Mountains may hold the answer. Trends in Ecology and Evolution, 15, 513–514.
- Körner, C. (2003) Alpine plant life. Springer Verlag, Berlin.
- Körner, C. (2007) The use of ‘altitude’ in ecological research. Trends in Ecology and Evolution, 22, 569–574.
- Lee, H.C., Chen, Y.J., Markhart, A.H. & Lin, T.Y. (2007) Temperature effects on systemic endoreduplication in orchid during floral development. Plant Science, 172, 588–595.
- Lomolino, M.V. (2001) Elevation gradients of species-density: historical and prospective views. Global Ecology and Biogeography, 10, 3–13.
- Manandhar, M.S. (1999) Evolution of Himalaya. Nepal nature’s paradise (ed. by T.C. Majupuria and R.K. Majupuria), pp. 13–17. M. Devi, Gwalior, India.
- Mani, A. (1981) The climate of the Himalaya. The Himalaya: aspects of change (ed. by J.S. Lall and A.D. Moddie), pp. 3–15. Oxford University Press, New Delhi.
- Mani, M.S. (1978) Ecology and phytogeography of high altitude plants of the north-west Himalaya. Oxford and IBH Publishing Co., New Delhi.
- McCain, C.M. (2005) Elevational gradients in diversity of small mammals. Ecology, 86, 366–372.
- McCain, C.M. (2007) Could temperature and water availability drive elevational species richness patterns? A global case study for bats. Global Ecology and Biogeography, 16, 1–13.
- McCullagh, P. & Nelder, J.A. (1989) Generalized linear models, 2nd edn. Chapman & Hall, London.
-
Miehe, G.,
Winniger, M.,
Boehner, J. &
Zand, Y.-L. (2001) Climate diagram map of high Asia.
Erdkunde, 55, 94–97.
10.3112/erdkunde.2001.01.06 Google Scholar
- Nogués-Bravo, D., Aráujo, M.B., Romdal, T. & Rahbek, C. (2008) Scale effects and human impact on the elevational species richness gradient. Nature, 453, 216–219.
- O’Brien, E.M. (1998) Water–energy dynamics, climate and prediction of woody plant species richness: an interim general model. Journal of Biogeography, 25, 379–398.
- O’Brien, E.M. (2006) Biological relativity to water–energy dynamics. Journal of Biogeography, 33, 1868–1888.
- Odland, A. & Birks, H.J.B. (1999) The altitudinal gradient of vascular plant richness in Aurland, western Norway. Ecography, 22, 548–566.
- Oommen, M.A. & Shankar, K. (2005) Elevational species richness patterns emerge from multiple local mechanisms in Himalaya woody plants. Ecology, 86, 3039–3047.
- Pearce, N.R. & Cribb, P.J. (2002) Flora of Bhutan: including a record of plants from Sikkim and Darjeeling, Vol. 3, Part 3. The orchids of Bhutan. Royal Botanical Garden, Edinburgh and Royal Government of Bhutan.
-
Petterson, B.D.,
Pacheco, V. &
Solari, S. (1996) Distribution of bats along an elevation gradient in the Andes of south-eastern Peru.
Journal of Zoology, 240, 637–658.
10.1111/j.1469-7998.1996.tb05313.x Google Scholar
- Press, J.K., Shrestha, K.K. & Sutton, D.A. (2000) Annotated checklist of the flowering plants of Nepal. Natural History Museum, London.
- R Development Core Team (2009) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available at: http://www.r-project.org
- Rahbek, C. (1995) The elevational gradient of species richness: a uniform pattern. Ecography, 18, 200–205.
- Rahbek, C. (1997) The relationship among area, elevation, and regional species richness in neotropical birds. The American Naturalist, 149, 875–902.
- Rahbek, C. (2005) The role of spatial scale and the perception of large-scale species-richness patterns. Ecology Letters, 8, 224–239.
- Rajbhandari, K.R. (1976) History of botanical explorations in Nepal. Journal of Bombay Natural History Society, 72, 468–481.
- Rajbhandari, K.R. & Dahal, S. (2004) Orchids of Nepal: a checklist. Botanica Orientalis, 4, 89–106.
- Rajbhandari, K.R., Bhattarai, S. & Joshi, R. (2000) Orchid diversity of Nepal and their conservation need. Proceedings of 8th International Workshop of BIO-REFOR: Biotechnology Applications for Reforestation and Biodiversity Conservation, Kathmandu, Nepal, 28 November–2 December, 1999 (ed. by M.S. Bista, R.B. Joshi, S.M. Amatya, A.V. Parajuli, M.K. Adhikari, H.K. Saiju, R. Thakur, K. Suzuki and K. Ishii), pp. 249–252. BIO-REFOR, IUFRO/SPDC, Tokyo.
-
Rosenzweig, M.L. (1995) Species diversity in space and time. Cambridge University Press, Cambridge.
10.1111/j.2006.0906-7590.04272.x Google Scholar
- Royal Botanic Gardens, Kew (2009) Monocot checklist. Available at: http://apps.kew.org/wcsp/qsearch.do (accessed 10 January 2009).
- Sanders, N.J., Lessard, J.-P., Fitzpatrick, M.C. & Dunn, R.R. (2007) Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains. Global Ecology and Biogeography, 16, 640–649.
-
Shakya, L.R. &
Shrestha, M.R. (2007) Two new taxa of Orchidaceae from central Nepal.
Edinburgh Journal of Botany, 64, 1–5.
10.1017/S0960428606000709 Google Scholar
- Shrestha, M.R., Shakya, L.R. & Ghimire, S.K. (2007) A new species of Malaxis Solander ex Swartz (Orchidaceae) from Dolpa, Nepal. Pleione, 1, 6–7.
- Shrestha, T.B. & Joshi, R.M. (1996) Rare, endemic and endangered plants in Nepal. WWF Nepal Programme, Kathmandu, Nepal.
- Singh, S.P. (2001) Central Himalaya biodiversity, ecosystem, characters and conservational approach. Environment and agriculture: biodiversity, agriculture and pollution in South Asia (ed. by S.R. Baral, P.K. Jha, S.B. Karmacharya, H.D. Lekhak, P. Lacoul and C.B. Baniya), pp. 15–18. Ecological Society, Kathmandu, Nepal.
- Stebbins, G.L. (1984) Polyploidy and distribution of the arctic-alpine flora: new evidence and new approaches. Botanica Helvetica, 94, 1–13.
- Terborgh, J. (1977) Bird species diversity on an Andean elevational gradient. Ecology, 58, 1007–1019.
- Vetaas, O.R. (2006) Biological relativity to water–energy dynamics: a potential unifying theory? Journal of Biogeography, 33, 1866–1867.
- Vetaas, O.R. & Ferrer-Castan, D. (2008) Richness of woody plants in the Iberian Peninsula: evaluating the importance of climate, topography and substrate at two spatial scales. Journal of Biogeography, 35, 1863–1878.
- Vetaas, O.R. & Grytnes, J.A. (2002) Distribution of vascular plant species richness and endemic richness along the Himalayan elevation gradient in Nepal. Global Ecology and Biogeography, 11, 291–301.
- Wang, W., Wang, Q., Li, S. & Wang, G. (2006) Distribution and species diversity of plant communities along transect on the northern Tibetan Plateau. Biodiversity and 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 and Distributions, 13, 845–854.
- Wangda, P. & Ohsawa, M. (2006) Gradational forest change along the climatically dry valley slopes of Bhutan in the midst of humid eastern Himalaya. Plant Ecology, 186, 109–128.
- White, K. & Sharma, B. (2000) Wild orchids in Nepal. White Lotus Press, Thailand.
- Whittaker, R.J., Willis, K.J. & Field, R. (2001) Scale and species richness: towards a general, hierarchical theory of species diversity. Journal of Biogeography, 28, 453–470.
- Winkel, G.V. (2006) Finding orchids in Nepal. The Orchid Review, 114, 188–191.
- Wolf, J.H.D. & Flamenco-S, A. (2003) Patterns in species richness and distribution of vascular epiphytes in Chiapas, Mexico. Journal of Biogeography, 30, 1689–1707.
- Yee, T.W. & Mitchell, N.D. (1991) Generalized additive models in plant ecology. Journal of Vegetation Science, 2, 587–602.