Allocating species in Grime’s strategy space: an alternative to trait-based approaches

Keywords: C-S-R theory, disturbance tolerance, functional attributes, Grime’s triangle, hydric stress tolerance.


Background:  The three primary-strategy theory proposed by Grime identifies stress and disturbance as key environmental factors leading to the emergence of distinct plant strategies. These are defined by a combination of stress and disturbance tolerance. Plant strategies are usually inferred from sets of traits, but this may lead to circular reasoning and artificial restrictions to species’ distribution in strategy space. 

Question: Can measurements of stress and disturbance tolerance be used to estimate the position of different species relative to each other in Grime’s strategy space?

Data description: Stress, disturbance, and abundances for 50 species at 25 0.5 ha sites.

Study site and dates: Semiarid grassland, Oaxaca, Mexico, 2014.

Methods: Species’ tolerance to stress and disturbance were inferred from abundances, and used to allocate species in Grime’s space. We tested if some attributes of our study species changed over the strategy space according to theoretical expectations.

Results: Most species were allocated towards high disturbance and low stress intensities. Species attributes were in line with the trends expected from their position in the strategy space. 

Discussion: Perhaps because of a long grazing history, most species were tolerant to disturbance. The allocation of species in the strategy space using stress and disturbance measurements seemed correct based on their attributes. Thus, our measurements seem to reflect the basic principles proposed by Grime. Our method provides relative positions in the strategy space, and (as previous work) requires defining somewhat arbitrary limits to such space if we wish to label species as ruderals, competitors or stress-tolerant.  


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Allocating species in Grime’s strategy space: an alternative to trait-based approaches


Adler D, Murdcoch D. 2014. rgl: 3D visualization device system (OpenGL). R package version 0.93.1098. (consulted: February 1, 2019)

Adler PB, Milchunas DG, Lauenroth WK, Sala OE, Burke IC. 2004. Functional traits of graminoids in semi-arid steppes: a test of grazing histories. Journal of Applied Ecology 41: 653-63. DOI:

Barba-Escoto L, Ponce-Mendoza A, García-Romero A, Calvillo-Medina RP. 2019. Plant community strategies responses to recent eruptions of Popocatépetl volcano, Mexico. Journal of Vegetation Sciences 30: 375-385. DOI:

Berryman AA. 2003. On principles, laws and theory in population ecology. Oikos 103: 695-701. DOI:

Bilton MC, Whitlock R, Grime JP, Marion G, Pakeman RJ. 2010. Intraspecific trait variation in grassland plant species reveals fine-scale strategy trade-offs and size differentiation that underpins performance in ecological communities. Botany 88: 939-952. DOI:

Bornhofen S, Barot S, Lattaud C. 2011. The evolution of CSR life-history strategies in a plant model with explicit physiology and architecture. Ecological Modelling 222: 1-10. DOI:

Briggs JM, Knapp AK. 1995. Interannual variability in primary production in tallgrass prairie: climate, soil water content, topographic position, and fire as determinants of aboveground biomass. American Journal of Botany 82: 1024-1030. DOI:

Campbell BD, Grime JP. 1992. An experimental test of plant strategy theory. Ecology 73: 15-29. DOI:

Chapin F, Schulze ED, Mooney HA. 1990. The ecology and economics of storage in plants. Annual Review of Ecology and Systematics 21: 423-447. DOI:

Cingolani AM, Renison D, Tecco PA, Gurvich DE, Cabido M. 2008. Predicting cover types in a mountain range with long evolutionary grazing history: a GIS approach. Journal of Biogeography 35: 538-551. DOI:

Chaves MM, Pereira JS, Maroco J, Rodrigues ML, Ricardo CPP, Osório ML, Carvalho I, Faria T, Pinheiro C. 2002. How plants cope with water stress in the field? Photosynthesis and growth. Annals of Botany 89: 907-916. DOI:

Díaz S, Noy-Meir I, Cabido M. 2001. Can grazing response of herbaceous plants be predicted from simple vegetative traits? Journal of Applied Ecology 38: 497-508. DOI:

Díaz S, Lavorel S, McIntyre S, Falczuk V, Casanoves F, Milchunas DG, Skapre C, Rusch G, Sternberg M, Noy-Meir I, Landsberg J, Zhang W, Clark H, Campbell BD. 2007. Plant trait responses to grazing - a global synthesis. Global Change Biology 13: 313-341. DOI:

Dornbush ME, Wilsey BJ. 2010. Experimental manipulation of soil depth alters species richness and co-occurrence in restored tallgrass prairie. Journal of Ecology 98: 117-125. DOI:

Fidelis A, Overbeck G, DePatta-Pillar V, Pfadenhauer, J. 2008. Effects of disturbance on population biology of the rosette species Eryngium horridum Malme in grasslands in southern Brazil. Plant Ecology 195: 55-67. DOI:

García HLA. 1996. La caprinocultura en la mixteca oaxaqueña: Orígenes. Ciencias 44: 28-31.

Grime JP. 1974. Vegetation classification by reference to strategies. Nature 250: 26-31. DOI:

Grime JP. 1977. Evidence for the existence of three primary strategies in plants and its relevance to ecological and evolutionary theory. American Naturalist 111: 1169-1194. DOI:

Grime JP. 1988. A comment on Loehle’s critique of the triangular model of primary plant strategies. Ecology 69: 1618-1620. DOI:

Grime JP. 2006. Plant Strategies, vegetation processes and ecosystem properties. Chichester: Wiley. ISBN: 978-0-470-85040-4

Gómez-García D, Azorín J, Aguirre AJ. 2009. Effects of small-scale disturbances and elevation on the morphology, phenology and reproduction of a successful geophyte. Journal of Plant Ecology 2: 13-20. DOI:

Haferkamp MR. 1988. Environmental factors affecting plant productivity. In: White RS, Short RE, eds. Achieving Efficient Use of Rangeland Resources. Miles City: Montana Agricultural Experimental Station, pp. 27-36.

Hanscom Z, Ting I. 1978. Responses of succulents to plant water stress. Plant Physiology 61: 327-330. DOI:

Hendrix SD, Trapp EJ. 1992. Population demography of Pastinaca sativa (Apiaceae): effects of seed mass on emergence, survival and recruitment. American Journal of Botany 79: 365-375. DOI:

Herben T, Klimešová J, Chytrý M. 2018. Philip Grime’s fourth corner: are there plant species adapted to high disturbance and low productivity? Oikos 127: 1125-1131. DOI:

Hobbs RJ, Huenneke LF. 1992. Disturbance, diversity, and invasion: Implications for conservation. Conservation Biology 6: 324-337. DOI:

Hodgson JG, Wilson PJ, Hunt R, Grime JP, Thompson K. 1999. Allocating C-S-R plant functional types: a soft approach to a hard problem. Oikos 85: 282-294. DOI:

Khurana E, Singh JS. 2004. Germination and seedling growth of five tree species from tropical dry forest in relation to water stress: impact of seed size. Journal of Tropical Ecology 20: 385-396. DOI:

Lavorel S, McIntyre S, Grigulis K. 1999. Plant response to disturbance in a Mediterranean grassland: How many functional groups? Journal of Vegetation Science 10: 661-672. DOI:

Li Y, Shipley B. 2017. An experimental test of CSR theory using a globally calibrated ordination method. PLOS ONE 12: e0175404. DOI:

Loehle C. 1988. Problems with the triangular model for representing plant strategies. Ecology 69: 284-286. DOI:

Louault F, Pillar VD, Aufrère J, Garnier E, Soussana JF. 2005. Plant traits and functional types in response to reduced disturbance in a semi-natural grassland. Journal of Vegetation Science 16: 151-160. DOI:

Martorell C, Peters EM. 2005. The measurement of chronic disturbance and its effects on the threatened cactus Mammillaria pectinifera. Biological Conservation 124: 199-207. DOI:

Martorell C, Martínez-López M. 2014. Informed dispersal in plants: Heterosperma pinnatum (Asteraceae) adjusts its dispersal mode to escape from competition and water stress. Oikos 123: 225-231. DOI:

Martorell C, Almanza-Celis CAI, Pérez-García EA, Sánchez-Ken JG. 2015. Co-existence in a species-rich grassland: competition, facilitation and niche structure over a soil depth gradient. Journal of Vegetation Science 26: 674-685. DOI:

McIntyre S, Lavorel S, Tremont RM. 1995. Plant life-history attributes: Their relationship to disturbance response in herbaceous vegetation. Journal of Ecology 83: 31-44. DOI:

McIntyre S, Lavorel S, Landsberg J, Forbes TDA. 1999. Disturbance response in vegetation: towards a global perspective on functional traits. Journal of Vegetation Science 10: 621-630. DOI:

Moles AT, Westoby M. 2006. Seed size and plant strategy across the whole life cycle. Oikos 113: 91-105. DOI:

Müller J, Heinze J, Joshi J, Boch S, Klaus VH, Fischer M, Prati D. 2014. Influence of experimental soil disturbances on the diversity of plants in agricultural grasslands. Journal of Plant Ecology 7: 509-517. DOI:

Ogburn RM, Edwards EJ. 2010. The ecological water-use strategies of succulent plants. Advances in Botanical Research 55:179-225. DOI:

Osmond CB, Austin MP, Berry JA, Billings WD, Boyer JS, Dacey JWH, Nobel PS, Smith SD, Winner WE. 1987. Stress physiology and the distribution of plants. BioScience 37: 38-48. DOI:

Pierce S, Brusa G, Vagge I, Cerabolini, BEL. 2013. Allocating CSR plant functional types: the use of leaf economics and size traits to classify woody and herbaceous vascular plants. Functional Ecology 27: 1002-1010. DOI:

Pierce S, Bottinelli A, Bassani I, Ceriani RM, Cerabolini BEL. 2014. How well do seed production traits correlate with leaf traits, whole-plant traits and plant ecological strategies? Plant Ecology 215: 1351-1359. DOI:

Pierce S, Negreiros D, Cerabolini BEL, Kattge J, Díaz S, Kleyer M, Shipley B, Wright SJ, Soudzilovskaia NA, Onipchenko VG, van Bodegom PM, Frenette-Dussault C, Weiher E, Pinho BX, Corneliseen JHC, Grime JP, Thompson K, Hunt R, Wilson PJ, Buffa G, Nyakunga OC, Reich PB, Caccianiga M, Mangili F, Ceriani RM, Luzzaro A, Brusa G, Siefert A, Barbosa NPU, Chapin III FS, Cornwell WK, Fang J, Fernandes GW, Garnier E, Le Stradic S, Peñuelas J, Melo FPL, Slaviero A, Tabarelli M, Tampucci D. 2017. A global method for calculating plant CSR ecological strategies applied across biomes world-wide. Functional Ecology 31: 444-457. DOI:

Quiroga RE, Golluscio RA, Blanco LJ, Fernández RJ. 2010. Aridity and grazing as convergent selective forces: an experiment with an Arid Chaco bunchgrass. Ecological Applications 20: 1876-1889. DOI:

R Core Team. 2017. R: A language and environment for statistical computing. R

Foundation for Statistical Computing, Vienna, Austria. (consulted: February 1, 2019)

Rajakaruna N, Boyd RS. 2008. Edaphic Factor. In: Jørgensen E, Fath, BD, eds. Encyclopedia of Ecology. Academic Press. 1201-1207. ISBN: 9780444637680

Raunkiaer C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press.

Robertson T, Atkins P. 2018. Essential vs. accidental properties. In: Zalta EN, ed. The Stanford Encylopedia of Philosophy Stanford: Metaphysics Research Lab.

Ruppert JC, Harmoney K, Henkin Z, Snyman HA, Sternberg M, Willims W, Linstädter A. 2015. Quantifying drylands’ drought resistance and recovery: the importance of drought intensity, dominant life history and grazing regime. Global Change Biology 21:1258-1270. DOI:

Schwinning S, Sala OE. 2004. Hierarchy of responses to resource pulses in arid and semi-arid ecosystems. Oecologia 141: 211-220. DOI:

Shea K, Chesson P. 2002. Community ecology theory as a framework for biological invasions. Trends in Ecology and Evolution 17: 170-176. DOI:

Sonnier G, Shipley B, Navas ML. 2010. Quantifying relationships between traits and explicitly measured gradients of stress and disturbance in early successional plant communities. Journal of Vegetation Science 21: 1014-1024. DOI:

Stanton ML, Roy BA, Thiede DA. 2000. Evolution in stressful environments. I. Phenotypic variability, phenotypic selection and response to selection in five distinct environmental stresses. Evolution 54: 93-111. DOI :[0093:EISEIP]2.0.CO;2

Tilman D. 1980. Resources: a graphical-mechanistic approach to competition and predation. The American Naturalist 116: 362-393. DOI:

Unger PW, Kaspar TC. 1994. Soil Compaction and Root Growth: A Review. Agronomy Journal 86: 759-766. DOI:

Villarreal-Barajas T, Martorell C. 2009. Species-specific disturbance tolerance, competition and positive interactions along an anthropogenic disturbance gradient. Journal of Vegetation Science 20: 1027-1040. DOI:

Van der Steen WJ, Scholten M. 1985. Methodological problems in evolutionary biology. IV. Stress and stress tolerance, an exercise in definitions. Acta Biotheoretica 34: 81-90. DOI:

Van der Steen WJ. 1993. Towards disciplinary disintegration in biology. Biology and Philosophy 8: 259-275. DOI:

Vibrans H. 2012. Malezas de México. Colegio de Posgraduados. (accessed February 18, 2019)

Wilson JB, Lee WG. 2000. C-S-R triangle theory: community-level predictions, tests, evaluation of criticism, and relation to other theories. Oikos 91: 77-96. DOI:

Wood SN. 2004. Stable and efficient multiple smoothing parameter estimation for generalized additive models. Journal of the American Statistical Association 99: 673-686. DOI:

Wulff RD. 1986. Seed size variation in Desmodium paniculatum: II. Effects on seedling growth and physiological performance. Journal of Ecology 74: 99-114. DOI:

Yuan ZY, Jiao F, Li YH, Kallenbach RL. 2016. Anthropogenic disturbances are key to maintain the biodiversity of grasslands. Scientific Reports 6: 22132. DOI:

How to Cite
Pedraza, F., & Martorell, C. (2019). Allocating species in Grime’s strategy space: an alternative to trait-based approaches. Botanical Sciences, 97(4), 649-660.