Elsa Cleland studies the responses of plant communities and ecosystems to global environmental changes, such as nitrogen deposition, elevated CO2, shifting precipitation, and invasive species. She is also interested in strategies for restoration of native plant communities in the context of present and future environmental changes. Her lab regularly measures carbon and other nutrients in samples collected both locally and globally.


Research in the Cleland lab focuses on the intersection between plant community and ecosystem ecology, utilizing a combination of experimental, observational and data-synthesis techniques. They have a particular focus on global changes such as climate change, altered precipitation, nitrogen deposition, and invasive species. Current research projects include: 1) Quantifying ecophysiological responses of California native shrubs to shifting precipitation and nitrogen availability in the greenhouse, in comparison to measurements made across natural gradients. 2) Identifying environmental manipulations that maximize the success of ecological restoration efforts in coastal sage scrub ecosystems, and 3) Evaluating the role of phenology (or timing) in structuring native and invasive plant communities. Research into the impacts of global change offers the opportunity to ask fundamental questions about how resources and climate influence the structure and functioning of natural ecosystems, while contributing to a greater understanding of these pressing environmental problems.


Select Publications

McKinney, J. G. & E. E. Cleland (2014). Root inputs influence soil water holding capacity and differentially influence the growth of native versus exotic annual species in an arid ecosystem. Restoration Ecology (in press) doi: 10.1111/rec.12139


Mauritz, M., E. Cleland, M. Merkley, & D. Lipson. (2014). The influence of altered rainfall regimes on early season N partitioning among early phenolgy annual plants, a late phenology shrub, and microbes in a semi-arid ecosystem.  Ecosystems (in press) doi: 10.1007/s10021-014-9800-6


Cleland, E.E., *E.E. Esch and *J.G. McKinney (2014). Priority effects vary with species identity and origin in an experiment varying the timing of seed arrival.  Oikos (in press) doi:10.1111/oik.01433


MacDougall, A. S., Bennett, J. R., Firn, J., Seabloom, E. W., Borer, E. T., Lind, E. M., Orrock, J. L., Harpole, W. S., Hautier, Y., Adler, P. B., Cleland, E., Davies, K., Melbourne, B., Prober, S. M., Bakker, J. D., Fay, P. A., Jin, V. L., Kendig, A., La Pierre, K. J., Moore, J., Morgan, J. and Stevens, C. J. (2014). Anthropogenic-based regional-scale factors most consistently explain plot-level exotic diversity in grasslands. Global Ecology and Biogeography. 23: 802-810


Wolkovich, E. M. & E.E. Cleland (2014). Phenological niches and the future of invaded ecosystems with climate change.  AoB PLANTS 6, plu013 (Editor’s Choice)


Borer, E. T., + 54 co-authors (2014). Herbivores and nutrients control grassland plant diversity via light limitation. Nature 508: 517–520.


Hallett,  L.M, J.S. Hsu, E. E. Cleland, S.L. Collins, T.L. Dickson, E. C. Farrer, L.A. Gherardi, K.L. Gross, R.J. Hobbs, L. Turnbull, K.N. Suding (2014). Biotic mechanisms of community stability shift along a precipitation gradient. Ecology 95:1693–1700.


Kopp, C. & E.E. Cleland. (2014). Shifts in elevational range limits and alpine plant species abundances observed over nearly five decades in a western North America mountain range. Journal of Vegetation Science 25: 135-146.


Seabloom, E., E. T. Borer, Y. Buckley, E. E. Cleland, + 69 co-authors. Predicting invasion in grassland ecosystems: is exotic dominance the real embarrassment of richness? (2013). Global Change Biology19:3677-3687.


Davies, T. J., ‡ E. M. Wolkovich, N. J. B. Kraft, N. Salamin, J. M. Allen, T. R. Ault, J. L. Betancourt, K. Bolmgren, E. E. Cleland, B. I. Cook, T. M. Crimmins, S. J. Mazer, G. J. McCabe, S. Pau, J. Regetz, M. D. Schwartz, & S. Travers (2013). Phylogenetic conservatism in plant phenology. Journal of Ecology  6:1520-1530.


Cleland, E. E., S. L. Collins, T. L. Dickson, E. C. Farrer, K. L. Gross, L. A. Gherardi, L. M. Hallett, R. J. Hobbs, J. S. Hsu, L. Turnbull & K. N. Suding. (2013). Sensitivity of grassland plant community composition to spatial versus temporal variation in precipitation. Ecology 94:1687-1696. link


Wainwright, C.W & E.E. Cleland (2013). Exotic species display greater germination plasticity and higher germination rates than native species across multiple cues. Biological Invasions 15: 2253-2264. link


Borer, E.T. M. E. S. Bracken, E. W. Seabloom, J. E. Smith, J. Cebrian, E. E. Cleland, J. J. Elser , W. F. Fagan, D. S. Gruner, W. S. Harpole, H. Hillebrand, A. J. Kerkhoff, and J. T. Ngai (2013). Global biogeography of autotroph chemistry: is insolation a driving force? Oikos 122:1121-1130.


Wolkovich, E. M.,  T. J. Davies, H. Schaefer, E. E. Cleland, B. I. Cook, S. E. Travers, C. G. Willis, C. C. Davis. (2013). Phenology and plant invasions: Climate change contributes to exotic species’ success in temperature-limited systems.  American Journal of Botany 100 :1407-1421.


O’Halloran, L, Borer, E., Seabloom, E., MacDougall, A.S., E. E. Cleland, R.L. McCulley, S. Hobbie, W. S. Harpole, N. M. DeCrappeo, C. Chu, J. D. Bakker, K. F. Davies, G. Du, J. Firn, N. Hagenah, K. S. Hofmockel, J. M. H. Knops, W. Li, B. A. Melbourne, J. W. Morgan, J. L. Orrock, S. M. Prober, and C. J. Stevens (2013). Regional contingencies in the relationship between aboveground biomass and litter in the world’s grasslands. PLoS ONE 8(2): e54988. doi:10.1371/journal.pone.0054988


Fraser, L.H. Henry, H.A.L., Carlyle, C.N., White, S.R., Beierkuhnlein, C., Cahill, J.F., Casper, B.B., Cleland, E., Collins, S.L., Dukes, J.S., Knapp, A.K., Lind, E., Long, R., Luo, Y. Reich, P.B., Smith, M.D., Sternberg, M. and Turkington, R.  (2013). Coordinated Distributed Experiments: an emerging tool for testing global hypotheses in ecology and environmental science. Frontiers in Ecology and the Environment 11:147-155.


Cleland, E.E., L.Larios & K.N. Suding (2013). Strengthening invasion filters to re-assemble native plant communities: soil resources and phenological overlap. Restoration Ecology. 21:390–398.   link


Cook, B., ‡E. Wolkovich, J. Davies, T. Ault, J. Betancourt, J. Allen, K. Bolmgren, E. Cleland, T. Crimmins, N. Kraft, L. Lancaster, S. Mazer, G. McCabe, B. McGill, C. Parmesan,S. Pau, J. Regetz, N. Salamin, M. Schwartz, &  S. Travers (2012). Sensitivity of spring phenology to warming across temporal and spatial climate gradients in two independent databases. Ecosystems. 15:1283-1294.


Cleland, E. E., J. M. Allen, T. M. Crimmins, J. A. Dunne, S. Pau, S. Travers, E. S. Zavaleta, and E. M. Wolkovich (2012). Phenological tracking enables positive species responses to climate change. Ecology93:1765–1771. link  (Research highlight in Nature Climate Change link)


Wainwright, C., ‡E.M. Wolkovich, & E. E. Cleland (2012). Seasonal priority effects: implications for invasion and restoration in a semi-arid system.  Journal of Applied Ecology 49:234-24. pdf (Reviewed on Faculty of 1000 link)


Wolkovich, E. M., B. I. Cook, J. M. Allen,T. M. Crimmins, J. L. Betancourt, S. Travers, S. Pau, J. Regetz, T. J. Davies, N. J. B. Kraft, T. R. Ault, K. Bolmgren, S. J. Mazer, G.J. McCabe, B. J. McGill, C. Parmesan, N. Salamin, M. D. Schwartz & E. E. Cleland (2012). Warming experiments underpredict plant phenological responses to climate change.  Nature 485:494-7 link (News & Views Forum link)


Gough, L., C.M. Clark, Cleland, E.E., S.L. Collins, J.E. Fargione, K.L. Gross, S.C. Pennings, & K.N. Suding (2012). Incorporating clonal growth form clarifies the role of plant height in response to nitrogen addition. Oecologia 169:1053-1062.


Cleland, E.E., C.M. Clark, S.L. Collins, J.E. Fargione, L. Gough, K.L. Gross, S.C. Pennings, & K.N. Suding (2011). Native and non-native species have different suites of traits: Evidence from a synthesis of nitrogen fertilization experiments. Journal of Ecology  99:1327-1338. link


Molina-Montenegro, E.E. Cleland, S. Watts M.A. & B. Broitman (2011).  Can a breakdown in competition – colonization tradeoffs help explain the success of exotic species in the California flora? Oikos 121:389-395.


Harpole, W.S., J.T. Ngai, E.E. Cleland, E.W. Seabloom, E.T. Borer, M.E.S. Bracken, J.J. Elser, D.S. Gruner, H. Hillebrand, J.B. Shurin, J.E. Smith (2011). Nutrient co-limitation of primary producer communities. Ecology Letters 14:852-862.


Wolkovich, E.M. & E.E. Cleland (2011).  The phenology of plant invasions: A community ecology perspecive.  Frontiers in Ecology and the Environment 9: 287–294. pdf  (Reviewed on Faculty of 1000 link)


Cleland, E.E. (2011). Trait divergence and the ecosystem impacts of invading species.  New Pytologist 189: 649-652. pdf


Cleland, E. E. (2011) Biodiversity and ecosystem stability. Nature Education Knowledge 2:2. link


Finzi, A.C, A. T. Austin, E. E. Cleland, S. D. Frey, B. Z. Houlton, & M. D. Wallenstein. (2011) Responses and feedbacks of coupled biogeochemical cycles to climate change: examples from terrestrial ecosystems. Frontiers in Ecology and the Environment.9:61-67.


Firn, J., J.L. Moore, A.S. MacDougall, E.T. Borer, E.W. Seabloom, J. HilleRisLambers, W.S. Harpole, E.E. Cleland, C.S. Brown, J.M H. Knops, S.M. Prober, D.A. Pyke, K.A. Farrell, J.D. Bakker, L.R. O’Halloran, P. B. Adler, S.L. Collins, C.M. D’Antonio, M.J. Crawley, E.M. Wolkovich, K.J. La Pierre, B.A. Melbourne, Y. Hautier, J.W. Morgan, A. D.B. Leakey, A. Kay, R. McCulley, K.F. Davies, C.J. Stevens, C.-J. Chu, K.D. Holl, J.A. Klein, P.A. Fay, N. Hagenah, K.P. Kirkman &Y.M. Buckley (2011). Abundance of introduced species at home predicts abundance away in herbaceous communities.  Ecology Letters 14:274-281.


Cadotte,M.W., E.T. Borer, J. Cavender-Bares, E. Cleland, K.F.. Davies, W. S.Harpole, E.W. Seabloom (2010). “Phylogenetic patterns differ for native and exotic plant communities across a richness gradient in Northern California.” Diversity and Distributions16: 892-901.


Sandel, B., L.J. Goldstein, N.J.B. Kraft, J.G. Okie, M.I. Shuldman, D. D. Ackerly, E.E. Cleland & K.N. Suding. (2010). Contrasting trait responses in plant communities to experimental and geographic variation in precipitation. New Phytologist 188:565–575.


Cleland, E.E. & W.S. Harpole (2010).  Nitrogen enrichment and plant communities. Annals of the New York Academy of Sciences 1195:46-61. pdf


Cadotte, M.W., T.J. Davies,J. Regetz,1 S.W. Kembel, E.E. Cleland & T. H. Oakley (2010). Phylogenetic diversity metrics for ecological communities: integrating species richness, abundance and evolutionary history. Ecology Letters 13: 96–105.


Hillebrand, H., E.T. Borer, M.E.S. Bracken, B.J. Cardinale, J. Cebrian, E.E. Cleland, J.J. Elser, D.S. Gruner, W.S. Harpole, J.T. Ngai, S. Sandin, E.W. Seabloom, J.B. Shurin, J.E. Smith & M.D. Smith (2009).  Herbivore metabolism and stoichiometry each constrain herbivory at different organizational scales across ecosystems. Ecology Letters 12:516-527.


Cleland, E.E., C.M. Clark, S.L. Collins, J.E. Fargione, L. Gough, K.L. Gross, D.G. Milchunas, S.C. Pennings, B. Bowman, I.C. Burke, W.K. Lauenroth, P. Robertson, J. Simpson, D. Tilman & K.N. Suding (2008). Species responses to nitrogen fertilization in herbaceous plant communities, and associated species traits.  Ecology 89:1175.  link


Collins, S.L., K.N. Suding, E.E. Cleland, M. Batty, S.C. Pennings, K.L. Gross, J.S. Grace, L. Gough, J.E. Fargione, and C.M. Clark (2008). Rank clocks and plant community dynamics. Ecology 89:3534-3541.


Chalcraft, D.R., S.B. Cox, C. Clark, E.E. Cleland, K.N. Suding, E. Weiher & D. Pennington (2008). Nitrogen enrichment studies often overestimate plant species loss at large spatial scales: the importance of beta diversity. Ecology 89:2165-2171.


Funk, J, E.E. Cleland, K.N. Suding & E.S. Zavaleta (2008). Restoration through re-assembly: plant traits and invasion resistance. TREE 23: 695-703.


Gruner, D.S., J.E. Smith, E.W. Seabloom, S.A. Sandin, J.T. Ngai, H. Hillebrand, W.S. Harpole, J.J. Elser, E.E. Cleland, M.E.S. Bracken, E.T. Borer & B.M. Bolker (2008). A cross-system synthesis of consumer and nutrient resource control on producer biomass.  Ecology Letters 11:740-755.


Drake, J.M., E.E. Cleland, M.C. Horner-Devine, E. Fleishman, C. Bowles, M.D. Smith, K. Carney, S. Emery, J. Gramling & D.B. Vandermast. (2008). Do Non-native plant species affect the shape of productivity-diversity relationships?  The American Midland Naturalist 159: 55–66.


Cleland E.E., Chuine I., Menzel A., Mooney H.A. & M.D. Schwartz (2007). Shifting plant phenology in response to global change. TREE 22:357-365. pdf


Clark, C.M., E.E. Cleland, S.L. Collins, J.E. Fargione, L. Gough, K.L. Gross, S.C. Pennings, K.N. Suding, & J.B. Grace (2007). Environmental and plant community determinants of species loss following nitrogen enrichment.  Ecology Letters 10: 596-607.


Elser, J.J., M.E. Bracken, E.E. Cleland, D.S. Gruner, W.S. Harpole, H. Hillebrand, J.T. Ngai, E.W. Seabloom, J.B. Shurin, & J.E. Smith (2007). Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater, marine, and terrestrial ecosystems.  Ecology Letters 10: 1115-1211.


Hillebrand, H., D.S. Gruner, M.E. Bracken, E.E. Cleland, J.J. Elser, W.S. Harpole, J.T. Ngai, E.W. Seabloom, J.B. Shurin, & J.E. Smith (2007).  Consumer versus resource control of producer diversity depends on ecosystem type and producer community structure. P.N.A.S. 104:10904-10909.


Peters, H.A., G. Hsu, E.E. Cleland, N.R. Chiariello, H.A. Mooney, & C.B. Field (2007). Responses of temporal distribution of gastropods to individual and combined effects of elevated CO2 and N deposition in annual grassland.  Acta Oecologica 31: 343-352.


Cleland E.E., N.R. Chiariello, S.R. Loarie, H.A. Mooney, & C.B. Field (2006). Diverse responses of

phenology to global changes in a grassland ecosystem. P.N.A.S. 103: 13740-13744. pdf


Cleland, E.E., H.A. Peters, H.A. Mooney, & C.B. Field (2006). Gastropod herbivory in response

to elevated CO2 and N deposition: impacts on plant community composition. Ecology 87:686–694 pdf


Peters, H.A., E.E. Cleland, H.A. Mooney, & C.B. Field (2006). Herbivore control of annual grassland composition in current and future environments. Ecology Letters 9: 86-94


Barnard, R., X. Le Roux, B. Hungate, E. Cleland, J. Blankinship, L. Barthes, P. Leadley (2006). Several components of global change alter nitrifying and denitrifying activities in an annual grassland. Functional Ecology 20:557-564.


Dukes, J.S., N.R. Chiariello, E.E. Cleland, L.A. Moore, M.R. Shaw, S. Thayer, T. Tobeck, H.A. Mooney & C.B. Field (2005). Responses of grassland production to single and multiple global environmental changes. P.L.o.S. Biology 3(10): e319


Henry, H.A.L, E.E. Cleland, C.B. Field & P.M. Vitousek (2005). Interactive effects of elevated CO2, N deposition and climate change on plant litter quality in a California annual grassland. Oecologia 142: 465-473


Pennings, S.C., C.M. Clark, E.E. Cleland, S.L. Collins, L. Gough, K.L. Gross, D.G. Milchunas & K.N. Suding (2005). Do individual plant species show predictable responses to nitrogen addition across multiple experiments? Oikos 110: 547-555


Suding, K.N., S.L. Collins, L. Gough, C.M. Clark, E.E. Cleland, K.L. Gross, D.G. Milchunas & S. Pennings (2005). Functional- and abundance-based mechanisms explain diversity loss due to N fertilization. P.N.A.S. 102: 4387-4392


Cleland, E.E., M.D. Smith, S.J. Andelman, C. Bowles, K.M. Carney, M.C. Horner-Devine, J. M. Drake, S. Emery & D.B. Vandermast (2004). “Invasion in space and time: Non-native species richness and relative abundance respond to interannual variation in productivity and diversity.” Ecology Letters 7:947-957 pdf


Hayhoe, K., D. Cayan, C.B. Field, P.C. Frumhoff, E.P. Maurer, N.L. Miller, S.C. Moser, S.H. Schneider, K.N. Cahill, E.E. Cleland, L. Dale, R. Drapek, R.M. Hanemann, L.S. Kalkstein, J. Lenihan, C.K. Lunch, R.P. Neilson, S.C. Sheridan, & J.H. Verville (2004). Emissions pathways, climate change, and impacts on California. P.N.A.S. 101:12422–12427


Zavaleta E.S., M.R. Shaw, N.R. Chiariello, B.D. Thomas, E.E. Cleland, C.B. Field, & H.A. Mooney (2003). Responses of a California annual grassland community to three years of warming, elevated CO2, precipitation increase, and N deposition. Ecological Monographs, 73:585-604


Field, C.B., M.R. Shaw, H.A. Mooney, E.S. Zavaleta, N.R. Chiariello, & E.E. Cleland (2002). Assessing environmental changes in grasslands: Response. Science 299:1844-1845


Shaw, R., E.S. Zavaleta, N.R. Chiarello, E.E. Cleland, H.A. Mooney, & C.B. Field (2002). Grassland responses to global environmental changes suppressed by elevated CO2. Science 298:1987-1990


Mooney, H.A., & E.E. Cleland (2001). The evolutionary impact of invasive species. P.N.A.S.  98:5446-5451


Novacek, M. J. & E.E. Cleland (2001). The current biodiversity extinction event: Scenarios for mitigation and recovery. P.N.A.S. 98:5466-6590


von Hippel, F. A., H. Frederick, & E.E. Cleland (2000). Population decline of the black and white colobus monkey (Colobus guereza) in the Kakamega Forest, Kenya. African Zoology, 35:69-75