Facilitation among plants in alpine environments in the face of climate change

Frontiers in Plant Science

Volumn 5, Issue AUG, 2014, Pages

  Anthelme, Fabien ^a,b   Cavieres, Lohengrin A ^c,d   Dangles, Olivier ^e,f  

^a INSTITUT DE RECHERCHE POUR LE DÉVELOPPEMENT   (France)

^b UNIVERSIDAD MAYOR DE SAN ANDRÉS   (Bolivia)

^c UNIVERSITY OF CONCEPCIÓN   (Chile)

^d INSTITUTO DE ECOLOGÍA Y BIODIVERSIDAD IEB   (Chile)

^e CNRS   (France)

^f UNIVERSITÉ PARIS SACLAY   (France)

Author keywords

Competition; Cushion plants; Early snowmelt; Facilitation; Global warming; Latitudinal gradient; Nurse plant; Stressgradient hypothesis

Indexed keywords

EID: 84907302689     PISSN: None     EISSN: 1664462X     Source Type: Journal    
DOI: 10.3389/fpls.2014.00387     Document Type: Review

References (131)

1. Adachi, N., Terashima, I., and Takahashi, M. (1996). Central die-back of monoclonal stands of Reynoutria japonica in an early stage of primary succession on Mount Fuji. Ann. Bot. 77, 477-486. doi: 10.1006/anbo.1996.0058
2. Adler, P. B., Dalgleish, H. J., and Ellner, S. P. (2012). Forecasting plant community impacts of climate variability and change: when do competitive interactions matter? J. Ecol. 100, 478-487. doi: 10.1111/j.1365-2745.2011.01930.x
3. Alatalo, J. M., and Little, C. J. (2014). Simulated global change: contrasting short and medium term growth and reproductive responses of a common alpine/Arctic cushion plant to experimental warming and nutrient enhancement. Springerplus 3:157. doi: 10.1186/2193-1801-3-157
4. Almeida, J. P., Montúfar, R., and Anthelme, F. (2013). Intraspecific trait variability reveals the specificities of successful altitudinal migration in a tropical alpine plant. Plant Ecol. Divers. 6, 423-433. doi: 10.1080/17550874.2012.702137
5. Anthelme, F., Buendia, B., Mazoyer, C., and Dangles, O. (2012). Unexpected mechanisms sustain the stress gradient hypothesis in a tropical alpine environment. J. Veg. Sci. 23, 62-72. doi: 10.1111/j.1654-1103.2011.01333.x
6. Anthelme, F., and Dangles, O. (2012). Plant-plant interactions in tropical alpine environments. Perspect. Plant Ecol. Evol. Syst. 14, 363-372. doi: 10.1016/j.ppees.2012.05.002
7. Araújo, M. B., and Luoto, M. (2007). The importance of biotic interactions for modelling species distributions under climate change. Global Ecol. Biogeogr. 16, 743-753. doi: 10.1111/j.1466-8238.2007.00359.x
8. Armas, C., Ordiales, R., and Pugnaire, F. I. (2004). Measuring plant interactions: a new comparative index. Ecology 85, 2682-2686. doi: 10.1890/03-0650
9. Armas, C., Schöb, C., and Gutiérrez, J. R. (2013). Modulating effects of ontogeny on the outcome of plant-plant interactions along stress gradients. New Phytol. 200, 7-9. doi: 10.1111/nph.12460
10. Aubert, S., Boucher, F., Lavergne, S., Renaud, J., and Choler, P. (2014). 1914-2014: a revised worldwide catalogue of cushion plants 100 years after Hauri and Schröter. Alp. Bot. 124, 59-70. doi: 10.1007/s00035-014-0127-x
11. Badano, E. I., and Marquet, P. A. (2009). Biogenic habitat creation affects biomass-diversity relationships in plant communities. Perspect. Plant Ecol. Evol. Syst. 11, 191-201. doi: 10.1016/j.ppees.2009.03.002
12. Bellard, C., Bertelsmeier, C., Leadley, P., Thuiller, W., and Courchamp, F. (2012). Impacts of climate change on the future of biodiversity. Ecol. Lett. 15, 365-377. doi: 10.1111/j.1461-0248.2011.01736.x
13. Benavides, J. C., Vitt, D. H., and Wieder, R. K. (2013). The influence of climate change on recent peat accumulation patterns of Distichia muscoides cushion bogs in the high-elevation tropical Andes of Colombia. J. Geophys. Res. Biogeo. 118, 1627-1635. doi: 10.1002/2013JG002419
14. Bertness, M., and Callaway, R. M. (1994). Positive interactions in communities. Trends Ecol. Evol. (Amst). 9, 191-193. doi: 10.1016/0169-5347(94)90088-4
15. Blois, J. L., Williams, J. W., Fitzpatrick, M. C., Jackson, S. T., and Ferrier, S. (2013). Space can substitute for time in predicting climate-change effects on biodiversity. Proc. Natl. Acad. Sci. U.S.A. 110, 9374-9379. doi: 10.1073/pnas.1220228110
16. Bobbink, R., Hornung, M., and Roelofs, J. G. M. (1998). The effects of air-borne nitrogen pollutants on species diversity in natural and semi-natural European vegetation. J. Ecol. 86, 717-738. doi: 10.1046/j.1365-2745.1998.8650717.x
17. Bradley, R. S., Vuille, M., Diaz, H. F., and Vergara, W. (2006). Threats to water supply in the tropical Andes. Science 312, 1755-1756. doi: 10.1126/science.1128087
18. Brooker, R. W. (2006). Plant-plant interactions and environmental change. New Phytol. 171, 271-284. doi: 10.1111/j.1469-8137.2006.01752.x
19. Brooker, R. W., and Callaghan, T. V. (1998). The balance between positive and negative plant interactions and its relationship to environmental gradients: a model. Oikos 81, 196-207. doi: 10.2307/3546481
20. Brooker, R. W., Maestre, F. E., Callaway, R. M., Lortie, C. J., Cavieres, L., Kunstler, G., et al. (2008). Facilitation in plant communities: the past, the present, and the future. J. Ecol. 96, 18-34.
21. Brooker, R. W., Travis, J. M., Clark, E. J., and Dytham, C. (2007). Modelling species' range shifts in a changing climate: the impacts of biotic interactions, dispersal distance and the rate of climate change. J. Theoretical Biol. 245, 59-65. doi: 10.1016/j.jtbi.2006.09.033
22. Caccianiga, M., Luzzaro, A., Pierce, S., Ceriani, R. M., and Cerabolini, B. (2006). The functional basis of a primary succession resolved by CSR classification. Oikos 112, 10-20. doi: 10.1111/j.0030-1299.2006.14107.x
23. Callaway, R. M. (2007). Positive Interactions and Interdependence in Plant Communities. Dordrechet: Springer.
24. Callaway, R. M., Brooker, R., Choler, P., Kikvidze, Z., Lortie, C. J., Michalet, R., et al. (2002). Positive interactions among alpine plants increase with stress. Nature 417, 844-848. doi: 10.1038/nature00812
25. Callaway, R. M., and Walker, L. R. (1997). Competition and facilitation: a synthetic approach to interactions in plant communities. Ecology 78, 1958-1965. doi: 10.1890/0012-9658(1997)078[1958:CAFASA]2.0.CO;2
26. Cannone, N., Sgorbati, S., and Guglielmin, M. (2007). Unexpected impacts of climate change on alpine vegetation. Front. Ecol. Environ. 5, 360-364. doi: 10.1890/1540-9295(2007)5[360:UIOCCO]2.0.CO;2
27. Cartenì, F., Marasco, A., Bonanomi, G., Mazzoleni, S., Rietkerk, M., and Giannino, F. (2012). Negative plant soil feedback explaining ring formation in clonal plants. J. Theoretical Biol. 313, 153-161. doi: 10.1016/j.jtbi.2012.08.008
28. Catorci, A., Cesaretti, S., Velasquez, J. L., and Zeballos, H. (2011). Plant-plant spatial interactions in the dry Puna (southern Peruvian Andes). Alp. Bot. 121, 113-121. doi: 10.1007/s00035-011-0097-1
29. Cauvy-Fraunié, S. (2010). Early Plant Succession on a Tropical Glacier Foreland in the Ecuadorian Andes. Paris: AgroParisTech (master report).
30. Cavieres, L. A., Arroyo, M., Peñaloza, A., Molina-Montenegro, M., and Torres, C. (2002). Nurse effect of Bolax gummifera cushion plants in the alpine vegetation of the Chilean Patagonian Andes. J. Veg. Sci. 13, 547-554.
31. Cavieres, L. A., and Badano, E. I. (2009). Do facilitative interactions increase species richness at the entire community level? J. Ecol. 97, 1181-1191. doi: 10.1111/j.1365-2745.2009.01579.x
32. Cavieres, L. A., Badano, E. I., Sierra-Almeida, A., Gómez, S., and Molina-Montenegro, M. A. (2006). Positive interactions between alpine plant species and the nurse cushion plant Laretia acaulis do not increase with elevation in the Andes of central Chile. New Phytol. 169, 59-69. doi: 10.1111/j.1469-8137.2005.01573.x
33. Cavieres, L. A., Brooker, R. W., Butterfield, B. J., Cook, B. J., Kikvidze, Z., Lortie, C. J., et al. (2014). Facilitative plant interactions and climate simultaneously drive alpine plant diversity. Ecol. Lett. 17, 193-202. doi: 10.1111/ele.12217
34. Cavieres, L. A., and Sierra-Almeida, A. (2012). Facilitative interactions do not wane with warming at high elevations in the Andes. Oecologia 170, 575-584. doi: 10.1007/s00442-012-2316-x
35. Cerabolini, B. E., Brusa, G., Ceriani, R. M., De Andreis, R., Luzzaro, A., and Pierce, S. (2010). Can CSR classification be generally applied outside Britain? Plant Ecol. 210, 253-261. doi: 10.1007/s11258-010-9753-6
36. Chapin, F. S., Walker, L. R., Fastie, C. L., and Sharman, L. C. (1994). Mechanisms of primary succession following deglaciation at Glacier Bay, Alaska. Ecol. Monogr. 64, 149-175. doi: 10.2307/2937039
37. Chen, I. C., Hill, J. K., Ohlemüller, R., Roy, D. B., and Thomas, C. D. (2011). Rapid range shifts of species associated with high levels of climate warming. Science 333, 1024-1026. doi: 10.1126/science.1206432
38. Choler, P., Michalet, R., and Callaway, R. M. (2001). Facilitation and competition on gradients in alpine plant communities. Ecology 82, 3295-3308. doi: 10.1890/0012-9658(2001)082[3295:FACOGI]2.0.CO;2
39. Cornelissen, J. H. C., Callaghan, T. V., Alatalo, J. M., Michelsen, A., Graglia, E., Hartley, A., et al. (2001). Global change and arctic ecosystems: is lichen decline a function of increases in vascular plant biomass? J. Ecol. 89, 984-994. doi: 10.1111/j.1365-2745.2001.00625.x
40. Couteron, P., Anthelme, F., Clerc, M., Escaff, D., Fernandez-Oto, C., and Tlidi, M. (2014). Plant clonal morphologies and spatial patterns as self-organized responses to resource-limited environments. P. Roy. Soc. A-Phys. Sci. (in press).
41. Crabtree, D., and Ellis, C. J. (2010). Species interaction and response to wind speed alter the impact of projected temperature change in a montane ecosystem. J. Veg. Sci. 21, 744-760.
42. Dangles, O., Herrera, M., and Anthelme, F. (2013). Experimental support of the stress-gradient hypothesis in herbivore-herbivore interactions. New Phytol. 197, 405-408. doi: 10.1111/nph.12080
43. Dormann, C. F., Van Der Wal, R., and Woodin, S. J. (2004). Neighbour identity modifies effects of elevated temperature on plant performance in the High Arctic. Global Change Biol. 10, 1587-1598. doi: 10.1111/j.1365-2486.2004.00830.x
44. Erschbamer, B., Kneringer, E., and Niederfriniger Schlag, R. (2001). Seed rain, soil seed bank, seedling recruitment, and survival of seedlings on a glacier foreland in the Central Alps. Flora 196, 304-312.
45. Erschbamer, B., Niederfriniger Schlag, R., and Winkler, E. (2008). Colonization processes on a central alpine glacier foreland. J. Veg. Sci. 19, 855-862. doi: 10.3170/2008-8-18464
46. Filazzola, A., and Lortie, C. J. (2014). A systematic review and conceptual frame-work for the mechanistic pathways of nurse plants. Global Ecol. Biogeogr. (in press).
47. Frénot, Y., Gloaguen, J. C., Cannavacciuolo, M., and Bellido, A. (1998). Primary succession on glacier forelands in the subantarctic Kerguelen Islands. J. Veg. Sci. 9, 75-84. doi: 10.2307/3237225
48. Garbarino, M., Lingua, E., Nagel, T. A., Godone, D., and Motta, R. (2010). Patterns of larch establishment following deglaciation of Ventina glacier, central Italian Alps. Forest Ecol. Manage. 259, 583-590. doi: 10.1016/j.foreco.2009.11.016
49. Gilman, S., Urban, M. C., Tewksbury, J., Gilchrist, G. W., and Holt, R. D. (2010). A framework for community interactions under climate change. Trends Ecol. Evol. 25, 325-331. doi: 10.1016/j.tree.2010.03.002
50. Gómez-Aparicio, L. (2009). The role of plant interactions in the restoration of degraded ecosystems: a meta-analysis across life-forms and ecosystems. J. Ecol. 97, 1202-1214. doi: 10.1111/j.1365-2745.2009.01573.x
51. Gottfried, M., Pauli, H., Futschik, A., Akhalkatsi, M., Barancok, P., Benito Alonso, J. L., et al. (2012). Continent-wide response of mountain vegetation to climate change. Nature Clim. Change 2, 111-115. doi: 10.1038/nclimate1329
52. Grabherr, G., Gottfried, M., and Pauli, H. (1994). Climate effects on mountain plants. Nature 369:448. doi: 10.1038/369448a0
53. Grassein, F., Lavorel, S., and Till-Bottraud, I. (2014). The importance of biotic interactions and local adaptation for plant response to environmental changes: field evidence along an elevational gradient. Global Change Biol. 20, 1462-1460. doi: 10.1111/gcb.12445
54. Grime, J. P. (1977). Evidence for the existence of three primary strategies in plants and its relevance to ecological and evolutionary theory. Am. Nat. 111, 1169-1194. doi: 10.1086/283244
55. Hättenschwiler, S., Handa, I. T., Egli, L., Asshoff, R., Ammann, W., and Körner, C. (2002). Atmospheric CO2 enrichment of alpine treeline conifers. New Phytol. 156, 363-375. doi: 10.1046/j.1469-8137.2002.00537.x
56. He, Q., and Bertness, M. D. (2014). Extreme stresses, niches and positive species interactions along stress gradients. Ecology 95, 1437-1443. doi: 10.1890/13-2226.1
57. He, Q., Bertness, M. D., and Altieri, A. H. (2013). Global shifts towards positive species interactions with increasing environmental stress. Ecol. Lett. 16, 695-706. doi: 10.1111/ele.12080
58. Hedberg, I., and Hedberg, O. (1979). Tropical-alpine life-forms of vascular plants. Oikos 33, 297-307. doi: 10.2307/3544006
59. Hobbie, S. E., Shevtsova, A., and Chapin, F. S. III. (1999). Plant responses to species removal and experimental warming in Alaskan tussock tundra. Oikos 84, 417-434. doi: 10.2307/3546421
60. Hu, Z., Anderson, N. J., Yang, X., and McGowan, S. (2014). Catchment-mediated atmospheric nitrogen deposition drives ecological change in two alpine lakes in SE Tibet. Global Change Biol. 20, 1614-1628. doi: 10.1111/gcb.12435
61. Hudson, J. M. G., Henry, G. H. R., and Cornwell, W. K. (2011). Taller and larger: shifts in Arctic tundra leaf traits after 16 years of experimental warming. Global Change Biol. 17, 1013-1021. doi: 10.1111/j.1365-2486.2010.02294.x
62. Hülber, K., Bardy, K., and Dullinger, S. (2011). Effects of snowmelt timing and competition on the performance of alpine snowbed plants. Perspect. Plant Ecol. Evol. Syst. 13, 15-26. doi: 10.1016/j.ppees.2011.01.001
63. IPCC. (2013). "Summary for policymakers" in Climate Change 2013: The Physical Science Basis Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change, eds T. F. Stocker, D. Qin, G.-K. Plattner, M. Tignor, S. K. Allen, J. Boschung, et al. (Cambridge: Cambridge University Press), 3-29.
64. Jentsch, A., Kreyling, J., and Beierkuhnlein, C. (2007). A new generation of climate-change experiments: events, not trends. Front. Ecol. Environ. 5, 365-374. doi: 10.1890/1540-9295(2007)5[365:ANGOCE]2.0.CO;2
65. Johnson, S. L., Kuske, C. R., Carney, T. D., Housman, D. C., Gallegos-Graves, L. V., and Belnap, J. (2012). Increased temperature and altered summer precipitation have differential effects on biological soil crusts in a dryland ecosystem. Global Change Biol. 18, 2583-2593. doi: 10.1111/j.1365-2486.2012.02709.x
66. Jones, G. A., and Henry, G. H. R. (2003). Primary plant succession on recently deglaciated terrain in the Canadian High Arctic. J. Biogeogr. 30, 277-296. doi: 10.1046/j.1365-2699.2003.00818.x
67. Kawai, T., and Tokeshi, M. (2007). Testing the facilitation-competition paradigm under the stress-gradient hypothesis: decoupling multiple stress factors. Proc. Biol. Sci. 274, 2503-2508. doi: 10.1098/rspb.2007.0871
68. Klanderud, K. (2005). Climate change effects on species interactions in an alpine plant community. J. Ecol. 93, 127-137. doi: 10.1111/j.1365-2745.2004.00944.x
69. Klanderud, K. (2008). Species-specific responses of an alpine plant community under simulated environmental change. J. Veg. Sci. 19, 363-372. doi: 10.3170/2008-8-18376
70. Klanderud, K., and Totland, O. (2005). The relative importance of neighbours and abiotic environmental conditions for population dynamic parameters of two alpine plant species. J. Ecol. 93, 493-501. doi: 10.1111/j.1365-2745.2005.01000.x
71. Klanderud, K., and Totland, O. (2007). The relative role of dispersal and local interactions for alpine plant community diversity under simulated climate warming. Oikos 116, 1279-1288. doi: 10.1111/j.0030-1299.2007.15906.x
72. Körner, C. (2003). Alpine Plant Life: Functional Plant Ecology of High Mountain Ecosystems. Berlin: Springer. doi: 10.1007/978-3-642-18970-8
73. Körner, C. (2007). The use of "altitude" in ecological research. Trends Ecol. Evol. 22, 569-574. doi: 10.1016/j.tree.2007.09.006
74. La Sorte, F. A., and Jetz, W. (2010). Projected range contractions of montane biodiversity under global warming. Proc. Biol. Sci. 277, 3401-3410. doi: 10.1098/rspb.2010.0612
75. Lavergne, S., Mouquet, N., Thuiller, W., and Ronce, O. (2010). Biodiversity and climate change: integrating evolutionary and ecological responses of species and communities. Annu. Rev. Ecol. Evol. Syst. 41, 321-350. doi: 10.1146/annurev-ecolsys-102209-144628
76. Lenoir, J., Gégout, J. C., Guisan, A., Vittoz, P., Wohlgemuth, T., Zimmermann, N. E., et al. (2010). Going against the flow: potential mechanisms for unexpected downslope range shifts in a warming climate. Ecography 33, 295-303.
77. Lenoir, J., Gégout, J. C., Marquet, P. A., de Ruffray, P., and Brisse, H. (2008). A significant upward shift in plant species optimum elevation during the 20th century. Science 320, 1768-1771. doi: 10.1126/science.1156831
78. le Roux, P. C., and McGeoch, M. A. (2004). The use of size as an estimator of age in the subantarctic cushion plant, Azorella selago (Apiaceae). Arctic Antarct. Alp. Res. 36, 509-517. doi: 10.1657/1523-0430(2004)36%5B509:TUOSAA%5D2.0.CO;2
79. le Roux, P. C., and McGeoch, M. A. (2008). Rapid range expansion and community reorganization in response to warming. Global Change Biol. 14, 2950-2962. doi: 10.1111/j.1365-2486.2008.01687.x
80. Le Roux, P. C., Shaw, J. D., and Chown, S. L. (2013). Ontogenetic shifts in plant interactions vary with environmental severity and affect population structure. New Phytol. 200, 241-250. doi: 10.1111/nph.12349
81. Loarie, S. R., Duffy, P. B., Hamilton, H., Asner, G. P., Field, C. B., and Ackerly, D. D. (2009). The velocity of climate change. Nature 462, 1052-1055. doi: 10.1038/nature08649
82. Lortie, C. J., Brooker, R. W., Choler, P., Kikvidze, Z., Michalet, R., Pugnaire, F. I., et al. (2004). Rethinking plant community theory. Oikos 107, 433-438. doi: 10.1111/j.0030-1299.2004.13250.x
83. Loza Herrera, S., Meneses, R. I., and Anthelme, F. (2014). Comunidades vegetales de los bofedales de la Cordillera Real bajo el calentamiento del clima. Revista de la Sociedad de Botanica de Bolivia. (in press).
84. Maestre, F. T., Callaway, R. M., Valladares, F., and Lortie, C. J. (2009). Refining the stress-gradient hypothesis for competition and facilitation in plant communities. J. Ecol. 97, 199-205. doi: 10.1111/j.1365-2745.2008.01476.x
85. Malkinson, D., and Tielbörger, K. (2010). What does the stress-gradient hypothesis predict? Resolving the discrepancies. Oikos 119, 1546-1552. doi: 10.1111/j.1600-0706.2010.18375.x
86. Marion, G. M., Henry, G. H. R., Freckman, D. W., Johnstone, J., Jones, G., Jones, M. H., et al. (1997). Open-top designs for manipulating field temperature in high-latitude ecosystems. Global Change Biol. 3, 20-32. doi: 10.1111/j.1365-2486.1997.gcb136.x
87. Martin, M., Gavazov, K., Koerner, C., Haettenschwiler, S., and Rixen, C. (2010). Reduced early growing season freezing resistance in alpine treeline plants under elevated atmospheric CO2. Global Change Biol. 16, 1057-1070. doi: 10.1111/j.1365-2486.2009.01987.x
88. Matthews, J. A. (1992). The Ecology of Recently Deglaciated Terrain A Geological Approach to Glacier Forelands and Primary Succession. Cambridge: Cambridge University Press.
89. McCluney, K. E., Belnap, J., Collins, S. L., González, A. L., Hagen, E. M., Nathaniel Holland, J., et al. (2012). Shifting species interactions in terrestrial dryland ecosystems under altered water availability and climate change. Biol. Rev. Camb. Philos. Soc. 87, 563-582. doi: 10.1111/j.1469-185X.2011.00209.x
90. McDonnell, T. C., Belyazid, S., Sullivan, T. J., Sverdrup, H., Bowman, W. D., and Porter, E. M. (2014). Modeled subalpine plant community response to climate change and atmospheric nitrogen deposition in Rocky Mountain National Park, USA. Environ. Pollut. 187, 55-64. doi: 10.1016/j.envpol.2013.12.021
91. Meier, E. S., Lischke, H., Schmatz, D. R., and Zimmermann, N. E. (2012). Climate, competition and connectivity affect future migration and ranges of European trees. Global Ecol. Biogeogr. 21, 164-178. doi: 10.1111/j.1466-8238.2011.00669.x
92. Michalet, R., Brooker, R. W., Cavieres, L. A., Kikvidze, Z., Lortie, C. J., and Pugnaire, F. I. (2006). Do biotic interactions shape both sides of the humped-back model of species richness in plant communities? Ecol. Lett. 9, 767-773. doi: 10.1111/j.1461-0248.2006.00935.x
93. Michalet, R., Schöb, C., Lortie, C. J., Brooker, R. W., and Callaway, R. M. (2014). Partitioning net interactions among plants along altitudinal gradients to study community responses to climate change. Functional. Ecol. 28, 75-86. doi: 10.1111/1365-2435.12136
94. Morris, W. F., and Doak, D. F. (1998). Life history of the long-lived gynodioecious cushion plant Silene acaulis (Caryophyllaceae), inferred from size-based population projection matrices. Am. J. Bot. 85, 784-793. doi: 10.2307/2446413
95. Murphy, J. M., Sexton, D. M., Barnett, D. N., Jones, G. S., Webb, M. J., Collins, M., et al. (2004). Quantification of modelling uncertainties in a large ensemble of climate change simulations. Nature 430, 768-772. doi: 10.1038/nature02771
96. Myers-Smith, I. H., Forbes, B. C., Wilmking, M., Hallinger, M., Lantz, T., Blok, D., et al. (2011). Shrub expansion in tundra ecosystems: dynamics, impacts and research priorities. Environ. Res. Lett. 6:045509. doi: 10.1088/1748-9326/6/4/045509
97. Nagy, L., and Grabherr, G. (2009). The Biology of Alpine Habitats. Oxford: Oxford University Press.
98. Niederfriniger Schlag, R., and Erschbamer, B. (2000). Germination and establishment of seedlings on a glacier foreland in the Central Alps, Austria. Arct. Antarct. Alp. Res. 32, 270-277. doi: 10.2307/1552525
99. Nogués-Bravo, D., Araújo, M. B., Errea, M. P., and Martinez-Rica, J. P. (2007). Exposure of global mountain systems to climate warming during the 21st Century. Global Environ. Change 17, 420-428. doi: 10.1016/j.gloenvcha.2006.11.007
100. Nyakatya, M. J., and McGeoch, M. A. (2008). Temperature variation across Marion Island associated with a keystone plant species (Azorella selago Hook.(Apiaceae). Polar Biol. 31, 139-151. doi: 10.1007/s00300-007-0341-8
101. Olsen, S. L., and Klanderud, K. (2014). Biotic interactions limit species richness in an alpine plant community, especially under experimental warming. Oikos 123, 71-78. doi: 10.1111/j.1600-0706.2013.00336.x
102. Pajunen, A. M., Oksanen, J., and Virtanen, R. (2011). Impact of shrub canopies on understorey vegetation in western Eurasian tundra. J. Veg. Sci. 22, 837-846. doi: 10.1111/j.1654-1103.2011.01285.x
103. Pauli, H., Gottfried, M., Dullinger, S., Abdaladze, O., Akhalkatsi, M., Alonso, J. L. B., et al. (2012). Recent plant diversity changes on Europe's mountain summits. Science 336, 353-355. doi: 10.1126/science.1219033
104. Poloczanska, E. S., Hawkins, S. J., Southward, A. J., and Burrows, M. T. (2008). Modeling the response of populations of competing species to climate change. Ecology 89, 3138-3149. doi: 10.1890/07-1169.1
105. Prévosto, B., Monnier, Y., Ripert, C., and Fernandez, C. (2012). To what extent do time, species identity and selected plant response variables influence woody plant interactions? J. Appl. Ecol. 49, 1344-1355. doi: 10.1111/jpe.12000
106. Rabatel, A., Francou, B., Soruco, A., Gomez, J., Caceres, B., Ceballos, J. L., et al. (2013). Review article of the current state of glaciers in the tropical Andes: a multi-century perspective on glacier evolution and climate change. Cryosphere Discuss. 7, 81-102. doi: 10.5194/tc-7-81-2013
107. Ralph, C. (1978). Observations on Azorella compacta (Umbelliferae), a tropical Andean cushion plant. Biotropica 10, 62-67. doi: 10.2307/2388107
108. Saccone, P., Delzon, S., Brun, J. J., and Michalet, R. (2009). The role of biotic interactions in altering tree seedling responses to an extreme climatic event. J. Veg. Sci. 20, 403-414. doi: 10.1111/j.1654-1103.2009.01012.x
109. Sattin, S. R., Cleveland, C. C., Hood, E., Reed, S. C., King, A. J., Schmidt, S. K., et al. (2009). Functional shifts in unvegetated, perhumid, recently-deglaciated soil do not correlate with shifts in soil bacterial community composition. J. Microbol. 47, 673-681. doi: 10.1007/s12275-009-0194-7
110. Scherrer, D., and Körner, C. (2010). Infra-red thermometry of alpine landscapes challenges climatic warming projections. Global Change Biol. 16, 2602-2613.
111. Scherrer, D., and Körner, C. (2011). Topographically controlled thermal-habitat differentiation buffers alpine plant diversity against climate warming. J. Biogeogr. 38, 406-416. doi: 10.1111/j.1365-2699.2010.02407.x
112. Schöb, C., Callaway, R. M., Anthelme, F., Brooker, R. W., Cavieres, L. A., Kikvidze, Z., et al. (2014a). The context-dependence of beneficiary feedback effects in plant facilitations. New Phytol. doi: 10.1111/nph.12908. [Epub ahead of print].
113. Schöb, C., Kammer, P. M., Choler, P., and Veit, H. (2009). Small-scale plant species distribution in snowbeds and its sensitivity to climate change. Plant Ecol. 200, 91-104. doi: 10.1007/s11258-008-9435-9
114. Schöb, C., Kammer, P. M., and Kikvidze, Z. (2012). Combining observational and experimental methods in plant-plant interaction research. Plant Ecol. Divers. 5, 27-36. doi: 10.1080/17550874.2012.674067
115. Schöb, C., Michalet, R., Cavieres, L. A., Pugnaire, F. I., Brooker, R. W., Butterfield, B. J., et al. (2014b). A global analysis of bi-directional interactions in alpine plant communities shows facilitators experiencing strong reciprocal fitness costs. New Phytol. 202, 95-105. doi: 10.1111/nph.12641
116. Searles, P. S., Flint, S. D., and Caldwell, M. M. (2001). A meta-analysis of plant field studies simulating stratospheric ozone depletion. Oecologia 127, 1-10. doi: 10.1007/s004420000592
117. Shevtsova, A., Haukioja, E., and Ojala, A. (1997). Growth response of subarctic dwarf shrubs, Empetrum nigrum and Vaccinium vitis-idaea, to manipulated environmental conditions and species removal. Oikos 84, 417-434.
118. Soliveres, S., and Maestre, F. (2014). Plant-plant interactions, environmental gradients and plant diversity: a global synthesis of community-level studies. Perspect. Plant Ecol. Evol. Syst. 16, 154-163. doi: 10.1016/j.ppees.2014.04.001
119. Stöcklin, J., and Bäumler, E. (1996). Seed rain, seedling establishment and clonal growth strategies on a glacier foreland. J. Veg. Sci. 9, 45-56. doi: 10.2307/3236415
120. Suding, K. N., Ashton, I. W., Bechtold, H., Bowman, W. D., Mobley, M. L., and Winkleman, R. (2008). Plant and microbe contribution to community resilience in a directionally changing environment. Ecol. Monogr. 78, 313-329. doi: 10.1890/07-1092.1
121. Sutherst, R. W., Maywald, G. F., and Bourne, A. S. (2007). Including species interactions in risk assessments for global change. Global Change Biol. 13, 1843-1859. doi: 10.1111/j.1365-2486.2007.01396.x
122. Thompson, L. G., Mosley-Thompson, E., Davis, M. E., and Brecher, H. H. (2011). Tropical glaciers, recorders and indicators of climate change, are disappearing globally. Ann. Glaciol. 52, 23-34. doi: 10.3189/172756411799096231
123. Tylianakis, J. M., Didham, R. K., Bascompte, J., and Wardle, D. A. (2008). Global change and species interactions in terrestrial ecosystems. Ecol. Lett. 11, 1351-1363. doi: 10.1111/j.1461-0248.2008.01250.x
124. Vittoz, P., Randin, C., Dutoit, A., Bonnet, F., and Hegg, O. (2009). Low impact of climate change on subalpine grasslands in the Swiss Northern Alps. Global Change Biol. 15, 209-220. doi: 10.1111/j.1365-2486.2008.01707.x
125. Walker, L. R., and del Moral, R. (2003). Primary Succession and Ecosystem Rehabilitation. Cambridge: Cambridge University Press. doi: 10.1017/CBO9780511615078
126. Walther, G. R. (2010). Community and ecosystem responses to recent climate change. Philos. Trans. R. Soc. Lond. B Biol. Sci. 365, 2019-2024. doi: 10.1098/rstb.2010.0021
127. Wheeler, J. A., Hoch, G., Cortés, A. J., Sedlacek, J., Wipf, S., and Rixen, C. (2014). Increased spring freezing vulnerability for alpine shrubs under early snowmelt. Oecologia 175, 219-229. doi: 10.1007/s00442-013-2872-8
128. Wipf, S., Gottfried, M., and Nagy, L. (2013). Climate change and extreme events-their impacts on alpine and arctic ecosystem structure and function. Plant Ecol. Divers. 6, 303-306. doi: 10.1080/17550874.2013.819533
129. Wipf, S., and Rixen, C. (2010). A review of snow manipulation experiments in Arctic and alpine tundra ecosystems. Polar Res. 29, 95-109. doi: 10.1111/j.1751-8369.2010.00153.x
130. Wipf, S., Rixen, C., and Mulder, C. P. (2006). Advanced snowmelt causes shift towards positive neighbour interactions in a subarctic tundra community. Global Change Biol. 12, 1496-1506. doi: 10.1111/j.1365-2486.2006.01185.x
131. Yang, Y., Niu, Y., Cavieres, L. A., and Hang, S. (2010). Positive associations between the cushion plant Arenaria polytrichoides (Caryophyllaceae) and other alpine plant species increase with altitude in the Sino-Himalayas. J. Veg. Sci. 21, 1048-1057. doi: 10.1111/j.1654-1103.2010.01215.x