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College of Forestry and Conservation
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Lab Publications

Articles Accepted / In review:

Nasto, M.K., S. Alvarez-Clare, Y. Lekberg, B.W. Sullivan, A.R. Townsend and C.C. Cleveland. Interactions among nitrogen fixation and soil phosphorus acquisition strategies in lowland tropical rain forests. Ecology Letters (submitted)

Knelman, J.E., S.K. Schmidt, H.J. Lynch, J.L. Darcy, S.C. Castle, C.C. Cleveland, D.R. Nemergut. Nutrient addition dramatically accelerates microbial community succession. PNAS (submitted)

Weintraub, S., S. Porder, C. Cleveland, A. Townsend. Topography controls soil nitrogen availability in a lowland tropical forest. Ecology (in revision)

Sullivan, B.W., W. K. Smith, A.R. Townsend, M.K. Nasto, S.C. Reed, R.L. Chazdon, C.C. Cleveland. A novel approach to quantify biological (N) fixation implies substantial human alteration of the tropical N cycle. PNAS (In press)

Taylor, P.G., W.R. Wieder, S. Weintraub, S. Cohen, C.C. Cleveland, A.R. Townsend. Organic forms dominate hydrologic nitrogen losses from a lowland tropical watershed. Ecology (in revision)

Published articles:


Sullivan, B.W., S.C. Castle, S.A. Alvarez Clare, S. Porder, S.C. Reed, L. Schreeg, A.R. Townsend and C.C. Cleveland. Assessing nutrient limitation in complex forested ecosystems: alternatives to large-scale fertilization experiments. Ecology 95: 668-681.

Taylor, P.,T. Bilinski, H. Fancher, C. Cleveland, D. Nemergut, S. Weintraub, W. Wieder and A. Townsend. 2014. Palm oil wastewater methane emissions and bioenergy potential. Nature Climate Change 4: 151-152.

Graham, E.B., W.R. Wieder, J.W. Leff, S.R. Weintraub, A.R. Townsend, C.C. Cleveland, L. Philippot and D.R. Nemergut. 2014. Do we need to understand microbial communities to predict ecosystem function? A comparison of statistical models of nitrogen cycling processes. Soil Biology and Biochemistry 68: 279-282.


Smith, W.K., C.C. Cleveland, S.C. Reed, S.W. Running.2013. Agricultural conversion without external water and nutrient inputs reduces biospheric vegetation productivity. Geophys. Res. Lett. 41, doi:10.1002/2013GL058857.

This article was also highlighted in Nature Geoscience

Cleveland, C.C., S.C. Reed, A.B. Keller, D.R. Nemergut, S.P. O'Neill, R. Ostertag and P.M. Vitousek. 2013. Litter quality versus microbial community controls over decomposition: A quantitative analysis. Oecologia 174: 283-294.

Cleveland, C.C., B.Z. Houlton, W.K. Smith, A.R. Marklein, S.C. Reed, W. Parton, S. Del Grosso, S.W. Running. 2013. Patterns of new versus recycled primary production in the terrestrial biosphere. Proceedings of the National Academy of Science of the USA 110: 12733-12737

Keville, M.P., S.C. Reed and C.C. Cleveland. 2013. Nitrogen cycling responses to mountain pine beetle disturbance in a high elevation whitebark pine ecosystem. PLoS ONE 8(6): e65004. doi:10.1371/journal.pone.0065004

Vitousek, P.M. D.N.L. Menge, S.C. Reed & C.C. Cleveland. 2013. Biological nitrogen fixation: Rates, patterns and ecological controls in terrestrial ecosystems. Philosophical Transactions of the Royal Society of London B 368: 20130119.

Reed, S. C., A. R. Townsend and C. C. Cleveland. 2013. Assessing the relationships among phosphorus, molybdenum and free-living nitrogen fixation in a tropical rain forest: Results from observational and experimental analyses. Biogeochemistry DOI 10.1007/s10533-013-9835-3

Ferrenberg, S., S. O'Neill, J. Knelman, B. Todd, S. Duggan, D. Bradley, T. Robinson, S.K. Schmidt, A.R. Townsend, M. Williams, C.C. Cleveland, B.A. Melbourne, L. Jiang and D.R. Nemergut. 2013. Changes in assembly processes in soil bacterial communities following a wildfire disturbance. ISME Journal. DOI: ISMEJ.2013.11


Weintraub, S.R., W.R. Wieder, C.C. Cleveland & A.R. Townsend.2012. Both absolute and relative soil resource availabilities drive exo-enzymatic shifts in a lowland tropical forest. Biogeochemistry DOI 10.1007/s10533-012-9812-2  

Keller, A.B., S.C. Reed, A.R. Townsend and C.C. Cleveland. Effects of canopy tree species on belowground biogeochemistry in a lowland wet tropical forest. Soil Biology and Biochemistry 58: 61-69.

Wieder, W.K., C.C. Cleveland, P.G. Taylor, D.R. Nemergut, E.Hinckley, L. Phillippot et al. Both increases and decreases in organic matter inputs reduce nitrate production and loss in lowland tropical forest. Biogeochemistry DOI 10.1007/s10533-012-9793-1

Schmidt, S.K., D.R. Nemergut, B. Todd, R.C. Lynch, J.L. Darcy, C.C. Cleveland and A.L. King. 2012. A simple method for determining limiting nutrients for photosynthetic crusts. Plant Ecology and Diversity DOI:10.1080/17550874.2012.738714

Cleveland, C.C. and B.W. Sullivan. 2012. Drought and tropical soil emissions. Nature 489: 211-212.

Reed, S.C., A.R. Townsend, E.A. Davidson and C.C. Cleveland. 2012. Patterns in foliar nutrient resorption and stoichiometry across multiple scales. New Phytologist 196: 173–180

Wickings, K., A.S. Grandy, S.C. Reed & C.C. Cleveland. 2012. The origin of litter chemical complexity during decomposition. Ecology Letters doi: 10.1111/j.1461-0248.2012.01837.x

Smith, W.K., C.C. Cleveland, S.R. Reed & S.W. Running. .2012. Quantification of bioenergy potential for the conterminous United States constrained by satellite observations of productivity. Environmental Science & Technology

Leff, J.W., W.R. Wieder, P.G. Taylor, A.R. Townsend, D.R. Nemergut, A.S. Grandy, & C.C. Cleveland. 2012. Experimental litterfall manipulation drives large and rapid changes in soil carbon cycling in a wet tropical forest. Global Change Biology doi: 10.1111/j.1365-2486.2012.02749.x (Biology Faculty of 1000 Selection)


Leff, J.W., D.R. Nemergut, A.S. Grandy, S.P. O'Neill, K. Wickings, A.R. Townsend and C.C. Cleveland. 2011. The effects of soil bacterial community structure on decomposition in a tropical rain forest. Ecosystems DOI: 10.1007/s10021-011-9510-2.

Knelman, J.E., T.M. Legg, S.P. O'Neill, C.L. Washenberger, A. Gonzalez, C.C. Cleveland & D.R. Nemergut. 2011. Bacterial community structure and function change in association with colonizer plants during early primary succession in a glacier forefield. Soil Biology & Biochemistry 46: 172-180.

Reed, S.C., C.C. Cleveland and A.R. Townsend. 2011.. Functional ecology of free-living nitrogen fixation: A contemporary perspective. Annual Review of Ecology, Evolution and Systematics 42: 489-512.

Cleveland, C.C. A.R. Townsend, P. Taylor, S. Alvarez-Clare, M.M.C. Bustamante, G. Chuyong, S.Z. Dobrowski, P. Grierson, K.E. Harms, B.Z. Houlton, A. Marklein, W. Parton, S. Porder, S.C. Reed, C.A. Sierra, W.L. Silver, E.V.J. Tanner & W.R. Wieder. 2011. Relationships among net primary productivity, nutrients and climate in tropical rain forest: A pan-tropical analysis. Ecology Letters 14: 939-947. (This paper also includes an Erratum)

Wieder, W.R., C.C. Cleveland and A.R. Townsend.2011. Throughfall exclusion and leaf litter manipulation drive higher rates of soil N2O emissions from a lowland wet tropical forest. Global Change Biology 17: 3195-3207.

Schmidt, S. K.,C. C. Cleveland, D. R. Nemergut, S. C. Reed, A. J. King and P. Sowell. 2011. Estimating phosphorus availability for microbial growth in an emerging landscape. Geoderma 163: 135-140.

Reed, S. C., P. M. Vitousek and C. C. Cleveland. 2011. Are patterns of nutrient limitation belowground consistent with those aboveground?: Results from a 4 million year chronosequence. Biogeochemistry 106: 323-336.

Wickings, K, A. S. Grandy, S. C. Reed and C. C. Cleveland. 2011. Management intensity effects on the biological and biochemical pathways of litter decomposition. Biogeochemistry 104: 365-379.


Townsend, A.R., C.C. Cleveland, B.Z. Houlton, C.B. Alden and J.W.C. White. 2010. Multi-element regulation of the tropical forest carbon cycle. Frontiers in Ecology and the Environment 9: 9-17.

Reed, S.C., A.R. Townsend, P.G. Taylor and C.C. Cleveland. 2010. Phosphorus Cycling in Tropical Forests Growing on Highly Weathered Soils. pp. 339-369 In: E Buenemann, A Oberson and E Frossard (Eds). Soil Biology 26: Phosphorus In Action – Biological Processes in Soil Phosphorus Cycling: Springer, Berlin.

Maron, J., M. Marler, J. Klironomos, C. Cleveland. 2010. Soil pathogens contribute to the positive plant diversity-productivity relationship. Ecology Letters 14: 36-41.

Nemergut, D. R., C. C. Cleveland, W. R. Wieder, C. L. Washenberger and A. R. Townsend. 2010. Plot-scale manipulations of organic matter inputs to soils correlate with shifts in microbial community composition in a lowland tropical rain forest. Soil Biology & Biochemistry 42: 2153-2160.

Reed, S. C., A. R. Townsend, C. C. Cleveland and D. R. Nemergut. 2010. Microbial community shifts influence patterns in tropical forest nitrogen fixation. Oecologia 164: 521-531.

Nemergut, D. R., E. K. Costello, M. Hamady, C. Lozupone, L. Jiang, S. K. Schmidt, N. Fierer, A. R. Townsend, C. C. Cleveland, L. Stanish and R. Knight. 2010. Global patterns in the biogeography of bacterial taxa. Environmental Microbiology 13: 135-144.

Cleveland, C. C., W. R. Wieder, S. C. Reed and A. R. Townsend. 2010. Experimental drought in a tropical rain forest increases soil carbon dioxide losses to the atmosphere. Ecology 91: 2313-2323.

Johnson, P. J. T., A. R. Townsend, C. C. Cleveland, P. M. Gilbert, R. W. Howarth, V. J. McKenzie, E. Reimankova and M. H. Ward. 2010. Linking environmental nutrient enrichment and disease emergence in humans and wildlife. Ecological Applications 20: 16-29.

Wieder, W. R., C. C. Cleveland and A. R. Townsend. 2010. Controls over leaf litter decomposition in wet tropical forests. Ecology 90:3333-3341.


Cleveland, C. C., B. Z. Houlton, C. Neill, S. C. Reed, A. R. Townsend and Y. Wang. 2009. Using indirect methods to constrain symbiotic nitrogen fixation rates: A case study from an Amazonian rain forest. Biogeochemistry 99: 1-13.

Fierer, N., M. S. Strickland, D. Liptzin, M. A. Bradford and C. C. Cleveland. 2009. Global patterns in belowground communities. Ecology Letters 12:1-12.

Sattin, S., C. C. Cleveland, E. Hood, S. C. Reed, A. J. King, S. K. Schmidt, M. S. Robeson, N. Ascarrunz and D. R. Nemergut. 2009. Functional shifts in unvegetated, perhumid, recently-deglaciated soils do not correlate with shifts in soil bacterial community composition. The Journal of Microbiology 47: 673-681.


Bowman, W. D., C. C. Cleveland, L. Halada, J. Hreško and J. S. Baron. Negative impact of nitrogen deposition on soil buffering capacity. 2008. Nature Geoscience doi:10.1038/ngeo339.

Townsend, A. R., G. P. Asner and C. C. Cleveland. 2008. The biogeochemical heterogeneity of tropical forests. Trends in Ecology & Evolution 23: 424-431.

Wieder, W. R., C. C. Cleveland and A. R. Townsend. 2008. Tropical tree species composition affects the oxidation of organic matter from litter. Biogeochemistry 88: 127-138

Reed, S. C., C. C. Cleveland and A. R. Townsend. 2008. Tree species control rates of free-living nitrogen fixation in a tropical rain forest. Ecology 89: 2924-2934.

Schmidt, S. K., S. C. Reed, D. R. Nemergut, S. Grandy, C. C. Cleveland, M. N. Weintraub, A. W. Hill, E. K. Costello, A. F. Meyer, A. M. Martin, J. Neff. 2008. The earliest stages of microbial and ecosystem succession in high-elevation, recently deglaciated soils. Proceedings of the Royal Society of London Series B: doi:10.1098/rspb.2008.0808


Cleveland, C. C. and D. Liptzin. 2007. C:N:P stoichiometry in soil: Is there a "Redfield ratio" for the microbial biomass? Biogeochemistry 85: 235-252 / DOI 10.1007/s10533-007-9132-0

D. R. Nemergut, S. P. Anderson, C. C. Cleveland, A. P. Martin, A. E. Miller, A. Seimon, S. K. Schmidt. 2007. Microbial community succession in unvegetated, recently-deglaciated soils. Microbial Ecology 53: 110-122.

Townsend, A. R., C. C. Cleveland, G. P. Asner and M. M. C. Bustamante. 2007. Controls over foliar N:P ratios in tropical rain forests. Ecology 88: 107-118.

Reed, S. C., C. C. Cleveland, A. R. Townsend. 2007. Controls over leaf litter and soil nitrogen fixation in two lowland tropical rain forests. Biotropica 39:585-592.

Schmidt, S. K., Costello, E. K., Nemergut, D. R., Cleveland, C. C., Reed, S. C., Weintraub, M. N., Meyer, A. F. and Martin, A. M. 2007. Microbial turnover and seasonal succession drive biogeochemical cycles in the alpine. Ecology 88: 1379-1385.


Cleveland, C. C. and A. R. Townsend. 2006. Nitrogen and phosphorus additions cause substantial losses of soil carbon from a lowland tropical rain forest. Proceedings of the National Academy of Sciences of the USA 103: 10316-10321.

Cleveland, C. C., D. R. Nemergut, S. K. Schmidt and A. R. Townsend. 2006. Increases in soil respiration following labile carbon additions linked to rapid shifts in soil microbial community composition. Biogeochemistry DOI 10.1007/s10533-006-9065-z

Cleveland, C. C., A. R. Townsend and S. K. Reed. 2006. Nutrient regulation of organic matter decomposition in a tropical rain forest. Ecology 87:492-503.


Cleveland, C. C., D. M. McKnight and A. R. Townsend. 2005. Trophic interactions: Links between the carbon and hydrologic cycles in terrestrial ecosystems. Pp. 1557-1574 in: M. G. Anderson and J. J. McDonnell (Eds). The Encyclopedia of Hydrological Sciences, Volume 3: John Wiley & Sons, West Sussex, UK (Invited contributor).

Galloway, J. N., G. Asner, E. W. Boyer, D. G. Capone, C. C. Cleveland, F. J. Dentener, P. Greene, E. Holland, R. W. Howarth, D. M. Karl, A. F. Michaels, S. P. Seitzinger, A. R. Townsend and C. Vorosmarty. 2004. Global and regional nitrogen cycles: Past, present and future. Biogeochemistry 70: 153-226.

Cleveland, C. C., A. R. Townsend, B. C. Constance, R. E. Ley, and S. K. Schmidt. 2004. Soil microbial dynamics in Costa Rica: Seasonal and biogeochemical constraints. Biotropica 36: 184-195.

Boyer, E. W., R. W. Howarth, J. N. Galloway, F. J. Dentener, P. Green, C. Vorosmarty, C. C. Cleveland and G. P. Asner. 2004. Nitrogen inputs to world regions. In A. R. Mosier, K. Syers and J. R. Freney (Eds.), Agriculture and the nitrogen cycle: assessing the impact of fertilizer use on food production and the environment. Washington, D.C., Island Press.

Cleveland, C. C., J. C. Neff, A. R. Townsend and E. Hood. 2004. Composition, dynamics and fate of leached dissolved organic matter in terrestrial ecosystems: Results of a decomposition experiment. Ecosystems 7: 275-285.

Bowman, W. D., H. Steltzer, T. N. Rosenstiel, C. C. Cleveland and C. L. Meier. 2004. Litter effects of two co-occurring alpine species on plant growth, microbial activity and immobilization of nitrogen. Oikos 104: 336-344.


Townsend, A. R., R. W. Howarth, F. A. Bazzaz, M. S. Booth, C. C. Cleveland, S. K. Collinge, A. P. Dobson, P. R. Epstein, E. A. Holland, D. R. Keeney, M. A. Malin, C. A. Rogers, P. Wayne and A. H. Wolfe. 2003. Human health effects of a changing global nitrogen cycle. Frontiers in Ecology and the Environment 1: 240-246.

Cleveland, C. C., A. R. Townsend, B. C. Constance and S. K. Schmidt. 2002. Soil microbial dynamics and biogeochemical cycling in lowland tropical rain forests and pastures of southwestern Costa Rica. Ecological Applications 13: 314-326.

Cleveland, C. C., A. R. Townsend and S. K. Schmidt. 2002. Phosphorus limitation of microbial processes in moist tropical forests. Ecosystems 5: 680-691.

Townsend, A. R., G. P. Asner, C. C. Cleveland, M. E. Lefer, and M. M. C. Bustamante. 2002. Unexpected changes in soil phosphorus dynamics following tropical deforestation to cattle pasture. Journal of Geophysical Research 107 (D20), 8067, doi: 10.1029/2001 JD000650, 2002.

Vitousek, P. M., K. Cassman, C. C. Cleveland, T. Crews, C. B. Field, N. B. Grimm, R. W. Howarth, R. Marino, L. Martinelli, E. B. Rastetter, and J. I. Sprent. 2002. Towards an ecological understanding of biological nitrogen fixation. Biogeochemistry 57/58: 1-45.

Asner, G. P., A. R. Townsend, W. Riley, P. A. Matson, J. C. Neff, and C. C. Cleveland. 2001. Physical and biogeochemical controls of terrestrial ecosystem responses to nitrogen deposition. Biogeochemistry 54: 1-39.

Cleveland, C. C., A.R. Townsend, D. S. Schimel, H. Fisher, R. W. Howarth, L. O. Hedin, S. S. Perakis, E. F. Latty, J. C. Von Fischer, A. Elseroad, and M. F. Wasson. 1999. Global patterns of terrestrial biological nitrogen (N2) fixation in natural ecosystems. Global Biogeochemical Cycles 13: 623-645.

Fahey, T. J., C. J. Williams, J. N. Rooney-Varga, C. C. Cleveland, K. M. Postek, S. D. Smith, and D. R. Bouldin. 1999. Nitrogen deposition in and around an intensive agricultural district in Central New York. Journal of Environmental Quality 28: 1585-1600.

Cleveland, C. C. and J. B. Yavitt. 1998. Microbial consumption of atmospheric isoprene in a temperate forest soil. Applied and Environmental Microbiology 64: 172-177.

Cleveland, C. C. and J. B. Yavitt. 1997. Consumption of atmospheric isoprene in soil. Geophysical Research Letters 24: 2379-2382.


Updated 2/2013

For questions about the research we do, or for information about opportunities in the lab, please contact me directly.

Cory Cleveland, Associate Professor
Department of Ecosystem & Conservation Sciences
College of Forestry & Conservation
CHCB 423/32 Campus Drive
University of Montana
Missoula, MT 59812

Phone: 406-243-6018 | Fax: 406-243-6656
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