Natural Resource Ecology Laboratory


Michael G. Ryan's Publications

Links lead to pdf files of Publications


Journal Articles (Updated 02/27/2015)

Asao S and MG Ryan.  2015. Carbohydrate regulation of photosynthesis and respiration from branch girdling in four species of wet tropical rain forest trees.  Tree Physiology, In Press, February 2015.

Atkin OK, KJ Bloomfield, PB Reich, MG Tjoelker et al. 2015. Global variability in leaf respiration in relation to climate, plant functional types and leaf traits. New Phytologist, In Press.

Quentin A, EA Pinkard, MG Ryan, DT Tissue et al. 2014.  Non-structural carbohydrates in woody plants cannot be quantitatively compared among laboratories. Tree Physiology, submitted November 2014.  Supplemental Material.

Asao S, R Bedoya-Arrieta and MG Ryan.  2015.  Variation in foliar respiration and wood CO2 efflux rates among species and canopy layers in a wet tropical forest.  Tree Physiology, In Press, November 2014.

Malone SL, CL Staudhammer, SF Oberbauer, P Olivas, MG Ryan, JL Schedlbauer, HW Loescher and G Starr.  2014.  El Niño Southern Oscillation (ENSO) enhances CO2 exchange rates in freshwater marsh ecosystems in the Florida Everglades. PLOS One DOI: 10.1371/journal.pone.0115058.

Malone SL, Staudhammer CL, Loescher HW, Olivas P, Oberbauer SF, Ryan MG, Schedlbauer J, Starr G. 2014. Seasonal patterns in energy partitioning of two freshwater marsh ecosystems in the Florida Everglades. Journal of Geophysical Research-Biogeosciences 119: 1487-1505.

Speckman HN, JM Frank, JB Bradford, BL Miles, WJ Massman, WJ Parton and MG Ryan.  2015.  Forest ecosystem respiration estimated from eddy covariance and chamber measurements under high turbulence and substantial tree mortality from bark beetlesGlobal Change Biology 21: 708–721, doi: 10.1111/gcb.12731.

Kelsey KC, KL Barnes, MG Ryan and JC Neff.  2014.  Short and long-term carbon balance of bioenergy electricity production fueled by forest treatments.  Carbon Balance and Management 9:6  doi:10.1186/s13021-014-0006-1. Also available from TreeSearch here.

White DA, JF McGrath, MG Ryan, M Battaglia, DS Mendham, J Kinal, GM Downes, DS Crombie and MA Hunt.  2014.  Managing for water-use efficient wood production in Eucalyptus globulus plantationsForest Ecology and Management 331: 272–280.

Ryan MG and S Asao.  2014.  Phloem transport in trees Tree Physiology 34: 1-4. 

Coble AP, A Autio, MA Cavaleri, D Binkley and MG Ryan.  2014.   Converging patterns of vertical variability in leaf morphology and nitrogen across seven Eucalyptus plantations in Brazil and Hawaii, USA. Trees – Structure and Function  28:1-15.

Ortega J, A Turnipseed, AB Guenther, TG Karl, DA Day, D Gochis, JA Huffman, AJ Prenni, EJT Levin, SM Kreidenweis, PJ DeMott, Y Tobo, EG Patton, A Hodzic, Y Cui, PC Harley, RH Hornbrook, EC Apel, RK Monson, ASD Eller, JP Greenberg, M Barth, P Campuzano-Jost, BB Palm, JL Jimenez, AC Aiken, MK Dubey, C Geron, J Offenberg, MG Ryan, PJ Fornwalt, SC Pryor, FN Keutsch, JP DiGangi, AWH Chan, AH Goldstein, GM Wolfe, S Kim, L Kaser, R Schnitzhofer, A Hansel, CA Cantrell, RL Mauldin III and JN Smith.  2014.  Overview of the Manitou Experimental Forest Observatory: site description and selected science results from 2008–2013.  Atmospheric Chemistry and Physics Discussions 14: 1647–1709;; doi:10.5194/acp-14-6345-2014.  Also available from TreeSearch here.

Ryan MG. 2013. Three decades of research at Flakaliden advancing whole-tree physiology, forest ecosystem and global change research.  Tree Physiology  33: 1123–1131.  doi:10.1093/treephys/tpt100.

Olivas PC, SF Oberbauer, DB Clark, DA Clark, MG Ryan, JJ O'Brien and H Ordonez. 2013. Comparison of direct and indirect methods for assessing leaf area index across a tropical rain forest landscape. Agricultural and Forest Meteorology 177: 110-116.

McDowell NG, MG Ryan,  MJB Zeppel and DT Tissue.  2013.  Improving our knowledge of drought-induced forest mortality through experiments, observations, and modelingNew Phytologist 200: 289-293. 

McDowell NG, RA Fisher, C Xu, JC Domec, T Hölttä, DS Mackay, JS Sperry, A Boutz, L Dickman, N Gehres, JM Limousin, A Macalady, J Martinez-Vilálta, M Mencuccini, JA Plaut, J Ogee, RE Pangle, DP Rasse, MG Ryan, S Sevanto, RH Waring, EA Yepez and WT Pockman.  2013.  Uncertainties and opportunities in modeling drought-induced vegetation mortalityNew Phytologist 200:304-321.

Malone SL, G Starr, CL Staudhammer and MG Ryan.  2013.  Effects of simulated drought on the carbon balance of Everglades short hydro-period marshGlobal Change Biology 19: 2511–2523, doi: 10.1111/gcb.12211.

Epron D, Y Nouvellon and MG Ryan.  2012.  Introduction to the invited issue on carbon allocation of trees and forestsTree Physiology 32: 639–643.

Rhoades CC, MA Battaglia, ME Rocca and MG Ryan.  2012.  Short- and medium-term effects of fuel reduction mulch treatments on soil nitrogen availability in Colorado conifer forestsForest Ecology and Management 276: 231–238.

Kashian DM, WH Romme, DB Tinker, MG Turner and MG Ryan. 2013.  Post-fire changes in forest carbon storage over a 300-year chronosequence of Pinus contorta-dominated forestsEcological Monographs 83: 49–66.

McKinley DC, Ryan MG, RA Birdsey, CP Giardina, ME Harmon, LS Heath, RA Houghton, RB Jackson, JF Morrison, BC Murray, DE Pataki and KE Skog. 2011.  A synthesis of current knowledge on forests and carbon storage in the United StatesEcological Applications 21: 1902-1924.

Westerling AL, MG Turner, EAH Smithwick, WH Romme and MG Ryan.  2011.  Continued warming could transform Greater Yellowstone fire regimes by mid-21st CenturyProceedings of the National Academy of Sciences 32: 13165-13170.

Conant RT, MG Ryan, GI Ågren, H Birge, EA Davidson, PE Eliasson, SE Evans, SD Frey, CP Giardina, F Hopkins, R Hyvönen, MUF Kirschbaum, JM Lavallee, J Leifeld, WJ Parton, JM Steinweg, MD Wallenstein, JÅM Wetterstedt and MA Bradford.  2011.  Temperature and soil organic matter decomposition rates – synthesis of current knowledge and a way forwardGlobal Change Biology 17: 3392–3404.

Ryan MG.  2011.  Tree responses to droughtTree Physiology 31: 237-239.

Battaglia MA, ME Rocca, CC Rhoades and MG Ryan.  2010.  Surface fuel loadings within mulching treatments in Colorado coniferous forests.  Forest Ecology and Management 260: 1557–1566.

Ryan MG.  2010.  Temperature and tree growthTree Physiology 30: 667–668.

Ryan MG, ME Harmon, RA Birdsey, CP Giardina, LS Heath, RA Houghton, RB Jackson, DC McKinley, JF Morrison, BC Murray, DE Pataki and KE Skog. 2010.  A Synthesis of the Science on Forests and Carbon for U.S. Forests. Ecological Society of America, Issues In Ecology 13: 1-16. The Issues in Ecology Series is available at the Ecological Society of America’s website here. Version in Español, translated by Patricia Inés Araujo and Amy T. Austin, IFEVA – CONICET, Facultad de Agronomía, Universidad de Buenos Aires aqui.

Ryan MG, JL Stape, D Binkley, S Fonseca, R Loos, EN Takahashi, CR Silva, S Silva, R Hakamada, JM Ferreira, AM Lima, JL Gava, FP Leite, G Silva, H Andrade and JM Alves. 2010. Factors controlling Eucalyptus productivity:  How water availability and stand structure alter production and carbon allocationForest Ecology and Management 259: 1695–1703.

Stape JL, D Binkley, MG Ryan, S Fonseca, R Loos, EN Takahashi, CR Silva, S Silva, R Hakamada, JM Ferreira, AM Lima, JL Gava, FP Leite, G Silva, H Andrade and JM Alves.  2010.  The Brazil Eucalyptus Potential Productivity Project:  Influence of water, nutrients and stand uniformity on wood productionForest Ecology and Management 259: 1684–1694.

Binkley D, JL Stape, WL Bauerle and MG Ryan.  2010.  Explaining growth of individual trees: Light interception and efficiency of light use by Eucalyptus at four sites in BrazilForest Ecology and Management 259: 1704–1713.

Hubbard RM, JL Stape, MG Ryan, AC Almeida and J Rojas.  2010.  Effects of irrigation on water use and water use efficiency in two fast growing Eucalyptus plantationsForest Ecology and Management 259: 1714–1721.

Ryan MG, MA Cavaleri, A Almeida de Campi, R Penchel, RS Senock and JL Stape.  2009.  Wood CO2 efflux and foliar respiration for Eucalyptus in Hawaii and BrazilTree Physiology 29: 1213–1222.

Bradford JB, P Weishampel, M-L Smith, R Kolka, RA Birdsey, SV Ollinger and MG Ryan.  2009.  Detrital carbon pools in temperate forests: magnitude and potential for landscape-scale assessment.  Canadian Journal of Forest Research 39: 802-813.

Cavaleri MA, SF Oberbauer, DB Clark, DA Clark and MG Ryan.  2010.  Height is the primary determinant of leaf morphology in a tropical rain forest canopyEcology 91: 1730–1739.

Bradford JB, P Weishampel, M-L Smith, R Kolka, RA Birdsey, SV Ollinger and MG Ryan.  2010.  Carbon pools and fluxes in small temperate forest landscapes: variability and implications for sampling design.   Forest Ecology and Management 259: 1245–1254.

Smithwick EAH, DM Kashian, MG Ryan and MG Turner.  2009.  Long-term ecosystem nitrogen storage and soil nitrogen availability in post-fire lodgepole pine ecosystemsEcosystems 12: 792-806.

Smithwick EAH, MG Ryan, DM Kashian, WH Romme, DB Tinker and MG Turner.  2009.  Modeling the effects of fire and climate change on carbon and nitrogen storage in lodgepole pine (Pinus contorta) stands.  Global Change Biology 15: 535-548.

Butnor JY, ML Pruyn, DC Shaw, ME Harmon, AN Mucciardi and MG Ryan.  2009.  Detecting defects in conifers with ground penetrating radarForest Pathology 39:309-322.

Bradford JB, RA Birdsey, LA Joyce and MG Ryan. 2008.  Tree age, disturbance history, and carbon stocks and fluxes in subalpine Rocky Mountain forestsGlobal Change Biology 14: 2882–2897.

Ryan MG, D Binkley and JL Stape.  2008.  Why don’t our stands grow even faster? Control of production and carbon cycling in eucalypt plantationsSouthern Forests 70: 99-104.

Sherrill KR, MA Lefsky, JB Bradford and MG Ryan 2008.  Forest structure estimation and pattern exploration from discrete return lidar in subalpine forests of the central RockiesCanadian Journal of Forest Research 38:2081-2096.

Cavaleri MA, SF Oberbauer and MG Ryan.  2008.  Foliar and ecosystem respiration in an old-growth tropical rain forestPlant Cell & Environment 31:473-483.

Clark DB, PC Olivas, SF Oberbauer, DA Clark and MG Ryan.  2008.  First direct landscape-scale measurement of tropical rain forest Leaf Area Index, a key driver of global primary productivityEcology Letters 11:163-172.

Stape JL, D Binkley and MG Ryan. 2008.  Production and carbon allocation in a clonal Eucalyptus plantation with water and nutrient manipulations.   Forest Ecology and Management 255: 920-930.

Litton CM, JW Raich and MG Ryan.  2007.  Carbon allocation in forest ecosystemsGlobal Change Biology 13: 2089-2109.

Hyvönen R, Ågren GI, Linder S, Persson T, Cotrufo MF, Ekblad A, Freeman M, Grelle A, Janssens IA, Jarvis PG, Kellomäki S, Lindroth A, Loustau D, Lundmark T, Norby RJ, Oren R, Pilegaard K, Ryan MG, Sigurdsson BD, Strömgren M, van Oijen M and Wallin G.  2007.  The likely impact of elevated [CO2], nitrogen deposition, increased temperature and management on carbon sequestration in temperate and boreal forest ecosystems: a literature review.  New Phytologist 173:463-480.

Palmroth S, R Oren, HR McCarthy, KH Johnsen, AC Finzi, JR Butnor, MG Ryan and WH Schlesinger.  2006. Aboveground sink strength in forests controls the allocation of carbon below ground and its [CO2]-induced enhancementProceedings of the National Academy of Sciences 130: 19362–19367.

Chapin FS, Woodwell GM, Randerson JT, Rastetter EB, Lovett GM, Baldocchi DD, Clark DA, Harmon ME, Schimel DS, Valentini R, Wirth C, Aber JD, Cole JJ, Goulden ML, Harden JW, Heimann M, Howarth RW, Matson PA, McGuire AD, Melillo JM, Mooney HA, Neff JC, Houghton RA, Pace ML, Ryan MG, Running SW, Sala OE, Schlesinger WH, Schulze ED.  2006.  Reconciling carbon-cycle concepts, terminology, and methods.  Ecosystems 9:1041-1050

Binkley D, Kashian DM, Boyden S, Kaye MW, Bradford JB, Arthur MA, Fornwalt PJ and Ryan MG.  2006.  Patterns of growth dominance in forests of the Rocky Mountains, USAForest Ecology and Management 236: 193–201.

Cavaleri MA, Oberbauer SF and MG Ryan.  2006.  Wood CO2 efflux in a primary tropical rain forestGlobal Change Biology 12:1-17.

Kashian DM, WH Romme, DB Tinker, MG Turner and MG Ryan.  2006.  Carbon storage on coniferous landscapes with stand-replacing firesBioScience 7:598-606.

Ryan MG, N Phillips and BJ Bond.  2006.  The hydraulic limitation hypothesis revisited. Plant, Cell and Environment 29: 367-381. ‘This is an electronic version of an article published in Plant, Cell and Environment: complete citation information for the final version of the paper, as published in the print edition of Plant, Cell and Environment, is available on the Blackwell Synergy online delivery service, accessible via the journal’s website at’

Binkley D, JL Stape, EN Takahashi and MG Ryan.  2006.  Tree-girdling to separate root and heterotrophic respiration in two Eucalyptus stands in Brazil.  Oecologia 148: 447-454.

Ryan MG and BE Law.  2005.  Interpreting, measuring, and modeling soil respiration. Biogeochemistry 73: 3-27.

Hubbard RM, MG Ryan, K Elder and CC Rhoades.  2005.  Seasonal patterns in soil surface CO2 flux under snow cover in 50 and 300 year old subalpine forests. Biogeochemistry 73: 93-107.

Binkley D, J Kaye, M Barry and MG Ryan. 2004.  First rotation changes in soil carbon and nitrogen in a Eucalyptus plantation in Hawaii. Soil Science Society of America Journal 68: 1713-1719

Harmon ME, K Bible, MG Ryan, DC Shaw, H Chen, J Klopatek and X Li. 2004. Production, respiration, and overall carbon balance in an old-growth Pseudotsuga-Tsuga forest ecosystem. Ecosystems 7: 98-512.

Ryan MG, D Binkley, JH Fownes, CP Giardina and RS Senock.  2004.  An experimental test of the causes of forest growth decline with stand age. Ecological Monographs 74: 393-414.

Pendall E, S Bridgham, PJ Hanson, B Hungate, DW Kicklighter, DW Johnson,  BE Law, YQ Luo, JP Megonigal, M Olsrud, MG Ryan and SQ Wan.  2004.   Below-ground process responses to elevated CO2 and temperature: a discussion of observations, measurement methods, and models.  New Phytologist 162: 311-322.

Giardina CP, D Binkley, MG Ryan, JH Fownes and RS Senock.  2004.  Belowground carbon cycling in a humid tropical forest decreases with fertilization. Oecologia 139: 545-550.

Stape JL, MG Ryan and D Binkley. 2004. Testing the 3-PG process-based model to simulate Eucalyptus growth with an objective approach to the soil fertility rating parameter. Forest Ecology and Management 193:219-234.

Stape JL, D Binkley and MG Ryan. 2004. Eucalyptus production and the supply, use and the efficiency of use of water, light and nitrogen across a geographic gradient in Brazil. Forest Ecology and Management 193:17-31.

Binkley D, JL Stape and MG Ryan. 2004. Thinking about efficiency of resource use in forests. Forest Ecology and Management 193:5-16.

Hubbard RM, MG Ryan, CP Giardina and HR Barnard. 2004. The effect of fertilization on sap flux and canopy conductance in a Eucalyptus saligna experimental forest. Global Change Biology 10:427-436.

Litton CM, MG Ryan and DH Knight. 2004. Effects of tree density and stand age on carbon allocation patterns in a postfire lodgepole pine ecosystem. Ecological Applications 14: 460-475.

Phillips N, BJ Bond, NG McDowell, MG Ryan and A Schauer. 2003. Leaf area compounds height-related hydraulic costs of water transport in Oregon white oak trees. Functional Ecology 17:832-840.

Giardina CP, MG Ryan, D Binkley and JH Fownes. 2003. Primary production and carbon allocation in relation to nutrient supply in an experimental tropical forest. Global Change Biology 9:1438-1450.

Litton CM, MG Ryan, DH Knight and PD Stahl. 2003. Soil-surface CO2 efflux and microbial biomass in relation to tree density thirteen years after a stand replacing fire in a lodgepole pine ecosystem. Global Change Biology 9:680-696.

Barnard HR and MG Ryan. 2003. A test of the hydraulic limitation hypothesis in fast-growing Eucalyptus saligna. Plant, Cell and Environment 26:1235-1245.

Phillips NG, MG Ryan, BJ Bond, NG McDowell, TM Hinckley and J Cermak. 2003. Reliance on stored water increases with tree size in three species in the Pacific Northwest. Tree Physiology 23: 237-245.

Litton CM, MG Ryan, DB Tinker and DH Knight. 2003. Below- and aboveground biomass in young post-fire lodgepole pine forests of contrasting tree density. Canadian Journal of Forest Research 33:351-363.

Ryan MG. 2002. Canopy processes research. Tree Physiology 22: 1035-1043.

McDowell N, N Phillips, C Lunch, BJ Bond and MG Ryan. 2002. An investigation of hydraulic limitation and compensation in large, old Douglas-fir trees. Tree Physiology 22:763-774.

McDowell N, H Barnard, BJ Bond, T Hinckley, RM Hubbard, H Ishii, B Köstner, FC Meinzer, JD Marshall, F Magnani, N Phillips, MG Ryan and D Whitehead. 2002. The relationship between tree height and leaf area:sapwood area ratio. Oecologia 132:12-20.

Giardina CP and MG Ryan. 2002. Total belowground carbon allocation in a fast growing Eucalyptus plantation estimated using a carbon balance approach. Ecosystems 5: 487-499.

Hubbard RM, BJ Bond, RS Senock and MG Ryan. 2002. The effect of branch height on leaf gas exchange, branch hydraulic conductance and branch sap flux in open grown ponderosa pine. Tree Physiology 22:575-581.

Vose JM and MG Ryan. 2002. Seasonal respiration of foliage, fine roots, and woody tissues in relation to growth, tissue N content, and photosynthesis. Global Change Biology 8:182-193.

Binkley D, JL Stape, MG Ryan, HR Barnard and JH Fownes. 2002. Age-related decline in forest ecosystem growth: an individual-tree, stand-structure hypothesis. Ecosystems 5:58-67.

Phillips N, BJ Bond, NG McDowell and MG Ryan. 2002. Canopy and hydraulic conductance in young, mature and old Douglas-fir trees. Tree Physiology 22: 205-211.

Williams M, BJ Bond and MG Ryan. 2001. Evaluating different soil and plant hydraulic constraints on tree function using a model and water flux data from ponderosa pine. Plant, Cell and Environment 24: 679-690.

Giardina CP, MG Ryan, RM Hubbard and D Binkley. 2001. Tree species and soil textural controls on carbon and nitrogen mineralization rates. Soil Science Society of America Journal 65:1272-1279.

Saxe H, MGR Cannell, Ø Johnsen, MG Ryan and G Vourlitis. 2001. Tree and forest functioning in response to global warming. New Phytologist 149:369-400.

Phillips N, Bond BJ and Ryan MG. 2001. Gas exchange and hydraulic properties in the crowns of two tree species in a Panamanian moist forest. Trees 15: 123-130.

Hubbard RM, V Stiller, MG Ryan and JS Sperry. 2001 Stomatal conductance and photosynthesis vary linearly with plant hydraulic conductance in ponderosa pine. Plant, Cell and Environment 24: 113-121.

Giardina CP and MG Ryan. 2000. Evidence that temperature does not influence the decomposition of organic carbon in forest mineral soils. Nature 404: 858-861.

Ryan MG, BJ Bond, BE Law, RM Hubbard, D Woodruff, E Cienciala and J Kucera. 2000. Transpiration and whole-tree conductance in ponderosa pine trees of different heights. Oecologia. 124: 553-560.

Bond BJ and MG Ryan. 2000. Comment on 'Hydraulic limitation of tree height: a critique' by Becker, Meinzer & Wullschleger. Functional Ecology 14: 135-140.

Brown PE, MG Ryan and TA Andrews. 2000. Historical surface fire frequency in ponderosa pine stands in Research Natural Areas, Central Rocky Mountains and Black Hills, USA. Natural Areas Journal 20: 133-139.

Hubbard RM, BJ Bond and MG Ryan. 1999. Evidence that hydraulic conductance limits photosynthesis in old Pinus ponderosa trees. Tree Physiology 19: 165-172.

Law BE, MG Ryan and PM Anthoni. 1999. Seasonal and annual respiration of a ponderosa pine ecosystem. Global Change Biology 5: 169-182.

Binkley D and MG Ryan. 1998. Net primary productivity and nutrient cycling in replicated stands of Eucalyptus saligna and Albizia facaltaria. Forest Ecology and Management 112: 79-85.

Cienciala E, J Kucera, MG Ryan and A Lindroth. 1998. Water flux in boreal forest during two hydrologically contrasting years; species regulation of canopy conductance and transpiration. Annales des Sciences Forestières 55: 47-61.

Ryan MG, D Binkley and JH Fownes. 1997. Age-related decline in forest productivity: pattern and process. Advances in Ecological Research 27: 213-262.

Ryan MG and BJ Yoder 1997.Hydraulic limits to tree height and tree growth. Bioscience 47: 235-242.

Margolis HA and MG Ryan. 1997. A physiological basis for biosphere—atmosphere interactions in the boreal forest: an overview. Tree Physiology 17:491-499.

Ryan MG, MB Lavigne and ST Gower. 1997. Annual carbon cost of autotrophic respiration in boreal forest ecosystems in relation to species and climate. Journal of Geophysical Research, BOREAS Special Issue, 102(D24): 28871-28884.

McCaughey JH, P Lafleur, D Joiner, D Jelinski and MG Ryan. 1997. Magnitudes and seasonal patterns of energy, water, and carbon exchanges at a boreal young jack pine forest in the BOREAS northern study area.  Journal of Geophysical Research, BOREAS Special Issue, 102(D24): 28997-29008.

Sellers PJ, FG Hall, RD Kelly, A Black, D Baldocchi, J Berry, MG Ryan, KJ Ranson, PM Crill, DP Lettenmaier, H Margolis, J Cihlar, J Newcomer, D Fitzjarrald, PG Jarvis, ST Gower, D Halliwell, D Williams, BGoodison, DE Wickland, and FE Guertin. 1997. BOREAS in 1997: Experiment Overview, Scientific Results and Future Directions.  Journal of Geophysical Research, BOREAS Special Issue, 102(D24): 28731-28770.

Lavigne MB, MG Ryan, DE Anderson, DD Baldocchi, PM Crill, DR Fitzjarrald, ML Goulden, ST Gower, JM Massheder, JH McCaughey, M Rayment and RG Striegl. 1997. Comparing nocturnal eddy covariance measurements to estimates of ecosystem respiration made by scaling chamber measurements. Journal of Geophysical Research, BOREAS Special Issue, 102(D24): 28977-28986.

Lavigne MB and MG Ryan. 1997. Growth and maintenance respiration rates of aspen, black spruce and jack pine stems at northern and southern BOREAS sites. Tree Physiology 17:543-551.

Ranne BM, WL Baker, T Andrews and MG Ryan. 1997. Natural variability of vegetation, soils, and physiography in the bristlecone pine forests of the Rocky Mountains. Great Basin Naturalist 57: 21-37.

Ryan MG, RM Hubbard, S Pongracic, RJ Raison and RE McMurtrie. 1996. Foliage, fine-root, woody-tissue and stand respiration in Pinus radiata in relation to nutrient content. Tree Physiology 16: 33-343.

Murty D, McMurtrie, RE and MG Ryan. 1996. Declining forest productivity in aging forest stands - a modeling analysis of alternative hypotheses. Tree Physiology 16: 187-200.

Ryan MG. 1995. Foliar maintenance respiration of subalpine and boreal trees and shrubs in relation to nitrogen content. Plant, Cell and Environment 18: 765-772.

Hubbard RM, MG Ryan and DL Lukens. 1995. A simple, battery operated temperature-controlled cuvette for respiration measurements. Tree Physiology 15: 175-179.

Sellers P, FG Hall, HA Margolis, R Kelly, D Baldocchi, J den Hartog, J Cilar, MG Ryan, BG Goodison, P Crill, J Ranson, D Lettenmaier and D Wickland. 1995. The Boreal Ecosystem-Atmosphere study (BOREAS): an overview and early results from the 1994 field year.  Bulletin of the American Meteorological Society 76: 1549-1577.

Ryan MG, ST Gower, RM Hubbard, RH Waring, HL Gholz, WP Cropper and SW Running. 1995. Stem maintenance respiration of four conifers in contrasting climates. Oecologia 101: 133-140.

Ryan MG, RM Hubbard, DL Clark and RL Sanford, Jr. 1994. Woody tissue respiration for Simarouba amara and Minquartia guianensis, two tropical wet forest trees with different growth habits. Oecologia 100: 213-220.

Ryan MG, S Linder, J Vose and RM Hubbard. 1994. Dark respiration in pines. Ecological Bulletins (Stockholm), 43: 50-63.

Yoder B, MG Ryan, RH Waring, AW Schoettle and MR Kaufmann. 1994. Evidence of reduced photosynthetic rates in old trees. Forest Science 40:513-527.

Waring RH, J Runyon, SN Goward, R McCreight, B Yoder and MG Ryan. 1993. Developing remote sensing techniques to estimate photosynthesis and annual forest growth across a steep climatic gradient in western Oregon, U.S.A. Studia Forestalia Suecica 194: 33-42.

Ryan MG and RH Waring. 1992. Stem maintenance and stand development in a subalpine lodgepole pine forest. Ecology 73: 2100-2108.

Binkley D, K Dunkin, D DeBell and MG Ryan. 1992. Litterfall cycling of nutrients in plantations of Eucalyptus and Albizia in Hawaii. Forest Science 38: 393-408.

Ryan MG. 1991. A simple method for estimating gross carbon budgets for vegetation in forest ecosystems. Tree Physiology 9: 255-266.

Oker-Blom P, MR Kaufmann and MG Ryan. 1991. Performance of a canopy light interception model for conifer shoots, trees, and stands. Tree Physiology 9: 227-243.

Rastetter EB, MG Ryan, GR Shaver, JM Melillo, KJ Nadelhoffer, JE Hobbie and JD Aber. 1991. A general biogeochemical model describing the responses of the C and N cycles in terrestrial ecosystems to changes in CO2, climate, and N deposition. Tree Physiology 9: 101-126.

Ryan MG. 1991. The effects of climate change on plant respiration. Ecological Applications 1: 157-167.

Melillo, J. M., T. V. Callaghan, F. I. Woodward, E. Salati, S. K. Sinha, J. Aber, V. Alexander, J. Anderson, A. Auclair, F. A. Bazzaz, A. Breymeyer, A. Clarke, C. B. Field, J. P. Grime, R. Gifford, J. Goudrian, R. Harris, I. Heaney, P. Holligan, P. J. Jarvis, L. A. Joyce, P. Levelle, S. Linder, A. Linkins, S. P. Long, A. E. Lugo, J. McCarthy, J. Morison, H. Nour, W. Oechel, M. Phillip, M. G. Ryan, D. Schimel, W. H. Schlesinger, G. R. Shaver, B. Strain, R. H. Waring, and M. Williamson.  1990.  Effects on Ecosystems.  Pages 284-310 in J. T. Houghton, G. J. Jenkins, and J. J. Ephraums, editors. Climate Change: The IPCC scientific assessment, Cambridge University Press, Cambridge, UK.

Ryan MG, JM Melillo and A Ricca. 1990. Comparing methods for determining proximate carbon fractions of forest litter. Canadian Journal of Forest Research 20: 166-171.

Ryan MG. 1990. Growth and maintenance respiration in stems of Pinus contorta and Picea engelmannii. Canadian Journal of Forest Research 20:48-57.

Ryan MG. 1989. Sapwood volume for three subalpine conifers: predictive equations and ecological implications. Canadian Journal of Forest Research 19: 1397-1401.

Schreuder HT, HG Li, MG Ryan and CT Scott. 1988. Adjusting estimates in two-way tables by incorporating outside information. Canadian Journal of Forest Research 18: 1280-1285.

Kaufmann MR and MG Ryan. 1986. Environmental, physiographic, and stand effects on individual tree growth in subalpine forests. Tree Physiology 2: 47-59.

Hinds TE and MG Ryan. 1985. Expansion of Sooty-bark and Ceratocystis canker expansion on aspen. Plant Disease 69:842-844.


Book Chapters

McDowell NG, BJ Bond, LT Dickman, MG Ryan, D Whitehead. 2011. Relationships between tree height and carbon isotope discrimination. Pages 255-286 in Meinzer FC, TE Dawson, and B Lachenbruch, editors, Size- and Age-Related Changes in Tree Structure and Function, Springer, New York, 510 p, ISBN 978-94-007-1241-6.

Bradford JB, P Weishampel, M-L Smith, RK Kolka, DY Hollinger, RA Birdsey, S Ollinger and MG Ryan. 2008. Landscape-Scale Carbon Sampling Strategy – Lessons Learned. Pages 227-238 in CM Hoover, editor, Field measurements for forest carbon monitoring: A landscape-scale approach, Springer, New York, NY, USA.

Bradford JB and MG Ryan. 2008. Quantifying soil respiration at landscape-scales. Pages 143-162 in CM Hoover, editor, Field measurements for forest carbon monitoring: A landscape-scale approach, Springer, New York, NY, USA.

CCSP, 2008: The effects of climate change on agriculture, land resources, water resources, and biodiversity. A Report by the U.S. Climate Change Science Program and the Subcommittee on Global Change Research. P Backlund, A Janetos, D Schimel, J Hatfield, K Boote, P Fay, L Hahn, C Izaurralde, BA Kimball, T Mader, J Morgan, D Ort, W Polley, A Thomson, D Wolfe, MG Ryan, SR Archer, RA Birdsey, CN Dahm, LS Heath, JA Hicke, DY Hollinger, TE Huxman, GS Okin, R Oren, JT Randerson, WH Schlesinger, D Lettenmaier, D Major, L Poff, S Running, L Hansen, D Inouye, BP Kelly, L Meyerson, B Peterson, R Shaw. US Environmental Protection Agency, Washington, DC, USA, 362 pp.

Giardina CP, MD Coleman, D Binkley, JE Hancock, JS King, EA Lilleskov, WM Loya, KS Pregitzer, MG Ryan, CC Trettin. 2005. The response of belowground carbon allocation in forests to global change. In D Binkley and O Menyailo (eds), 119-154. Tree species effects on soils: implications for global change. NATO Science Series, Kluwer Academic Publishers, Dordrecht.

Ryan MG, ER Hunt Jr., RE McMurtrie, GI Ågren, JD Aber, AD Friend, EB Rastetter, WJ Pulliam, RJ Raison and S Linder. 1996. Comparing models of ecosystem function for coniferous forests. I. Model description and validation. Pages 313-362 in AI Breymeyer, DO Hall, GI Ågren and JM Melillo, editors, Global Change: Effects on Coniferous Forests and Grasslands (SCOPE), John Wiley and Sons.

Ryan MG, RE McMurtrie, GI Ågren, Hunt, ER, Jr., JD Aber, AD Friend, EB Rastetter and WJ Pulliam. 1996. Comparing models of ecosystem function for coniferous forests. II. Predictions of response to changes in atmospheric CO2 and climate. Pages 363-387 in AI Breymeyer, DO Hall, GI Ågren and JM Melillo, editors, Global Change: Effects on Coniferous Forests and Grasslands (SCOPE), John Wiley and Sons.

Sprugel DF, MG Ryan, JR Brooks, KA Vogt and TA Martin. 1994. Respiration from the organ level to the stand. Pages 255-299 in WK Smith and TM Hinckley, editors, Resource Physiology of Conifers: Acquisition, Allocation, and Utilization, Academic Press, Boca Raton, Florida, USA.

Waring RH and MG Ryan. 1995. Carbon balance Modelling. Pages 480-491 in A Bisio and SG Boots, editors, Encyclopedia of Energy Technology and the Environment, 4th Edition, John Wiley and Sons, New York, NY, USA.


Other Publications

Ryan, MG. 2012. Disturbance Effects on Forest Carbon Dynamics. In Swanston C, MJ Furniss, K Schmitt, J Guntle, J Jeffrey, M Janowiak, S Hines, eds. 2012. Forest and grassland carbon in North America: A short course for land managers. Gen. Tech Rep. NRS- 93. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northern Research Station. [DVD].

Smithwick EAH, Westerling AL, MG Turner, WH Romme and MG Ryan. 2011. Vulnerability of Landscape Carbon Fluxes to Future Climate and Fire in the Greater Yellowstone Ecosystem. In Questioning Greater Yellowstone’s Future: Climate, Land Use, and Invasive Species. Proceedings of the 10th Biennial Scientific Conference on the Greater Yellowstone Ecosystem. October 11–13, 2010, Mammoth Hot Springs Hotel, Yellowstone National Park. C. Andersen, ed. Yellowstone Center for Resources (Yellowstone National Park, WY) and University of Wyoming William D. Ruckelshaus Institute of Environment and Natural Resources (Laramie, WY).

Battaglia MA, K Nelson, DM Kashian, and MG Ryan. 2010. Forest biomass and tree planting for fossil fuel offsets in the Colorado Front Range. Pages 81-93 in Jain TB, RT Graham and J Sandquist, technical editors. Integrated management of carbon sequestration and biomass utilization opportunities in a changing climate: Proceedings of the 2009 National Silviculture Workshop; 2009 June 15-18; Boise, ID. Proceedings RMRS-P-61. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 351 p.

Binkley D, J-P Laclau, JL Stape and MG Ryan. 2010. Applying ecological insights to increase productivity in tropical plantations. Forest Ecology and Management. 259: 1681–1683.

Ryan MG. 2009. What Is the Role of Forests in the U.S. Carbon Balance? In Furniss MJ, CI Millar, DL Peterson, LA Joyce, RP Neilson and JE Halofsky, editors. Adapting to climate change: a short course for land managers. General Technical Report PNW-GTR-789. Portland, OR: US Department of Agriculture, Forest Service, Pacific Northwest Research Station. DVD and online at

Ryan MG, DM Kashian, EAH Smithwick, WH Romme, MG Turner and DB Tinker. 2008. Final Report, Carbon cycling at the landscape scale: the effect of changes in climate and fire frequency on age distribution, stand structure, and net ecosystem production. JFSP Project Number: 03-1-1-06

Ryan, MG. 2008. Forests and Carbon Storage. (June 04, 2008). U.S. Department of Agriculture, Forest Service, Climate Change Resource Center.

Nelson KN, DM Kashian and MG Ryan. 2007. Planting burned areas for carbon sequestration. Final Report, Coalition for the Upper South Platte.

Nelson KN, DM Kashian and MG Ryan. 2007. Forest biomass removals for fossil fuels offsets. Final Report, Coalition for the Upper South Platte.

Campbell RE and MG Ryan. 1982. Precipitation and temperature characteristics of forested watersheds in north-central Arizona. USDA Forest Service, General Technical Report RM-93, 12p.

Rogers, JJ, JM Prosser, LD Garrett and MG Ryan. 1984. ECOSIM: a system for projecting multiresource outputs under alternative forest management regimes. RM Administrative Report.

Ryan, MG and WW Covington. 1986. Effects of prescribed burning in ponderosa pine on soil inorganic nitrogen content. USDA Forest Service Research Note, RM-464, 5p.

Kaufmann, MR, CA Troendle, MG Ryan and HT Mowrer. 1987. Trees--the link between silviculture and hydrology. Symposium proceedings, Management of Subalpine Forests: Building on 50 Years of Research. USDA Forest Service, General Technical Report, RM-149, p. 54-60.

Ryan, MG. 1987. Growth efficiency, leaf area and sapwood volume in subalpine conifers. Symposium proceedings, Management of Subalpine Forests: Building on 50 Years of Research. USDA Forest Service, General Technical Report, RM-149, p. 180-184.

Schuler, CE, DC Markstrom and MG Ryan. 1989. Fibril angle in young-growth ponderosa pine as related to site index, d.b.h, and location in tree. USDA Forest Service Research Note, RM-492, 4p.

Ryan, MG, LA Joyce, T Andrews and K Jones. 1994. Research Natural Areas of the USDA Forest Service in Colorado, Wyoming, Nebraska, North Dakota, and South Dakota. USDA Forest Service General Technical Report, RM-251, 58 p.