Publications 2018-01-08T16:21:40+00:00


Lab affiliation: *undergraduate student, **graduate student, †post doctoral fellow, ‡research staff

In Press

Blume-Werry G**, Lindén E, Andresen L, Classen AT, Sanders NJ, von Oppen J, Sundqvist MK. (in press) Proportion of fine roots, but not plant biomass allocation belowground, increases with elevation in arctic tundra heath communities. Journal of Vegetation Science.

Liu H, Zhaorong M, Lin L, Wang Y, Zhang Z, Zhang F, Wang H, Liu L, Zhu B, Cao G, Zhao X, Sanders NJ, Classen AT, Reich PB, He JS (2018) Shifting plant species composition in response to climate change stabilizes grassland primary production. PNAS 

Read QD**, Henning JA**, Classen AT, Sanders NJ (2018) Aboveground resilience to species loss but belowground resistance to nitrogen addition in montane plant communities. Journal of Plant Ecology View online.


Niu S, Classen AT, Luo Y (2018) Functional traits along a transect. Functional Ecology 32:4-9. View online.


Classen AT, Inouye BD (2017) Editorial. Ecological Monographs. 87:3. View online.

Bradford MA, Veen GF, Bonis A, Bradford EM, Classen AT, Cornelissen JHC, Crowther TW, De Long JR, Kardol P, Manrubia-Freixa M, Freshet GT, Maynard DS, Newman G‡, van Logtestijn RSP, Viketoft M, Wardle DA, Wieder WR, Wood SA, van der Putten WH (2017) Testing the hierarchical model of litter decomposition. Nature Ecology & Evolution doi:10.1038/s41559-017-0367-4 View online.

Hendershot JN*, Read QD**, Henning JA**, Sanders NJ, Classen AT (2017) Consistently inconsistent drivers of patterns of microbial diversity and abundance at macroecological scales. Ecology 98:1757-1763. View online.

Jiang J†, Moore JAM**, Priyadarshi A, Classen AT (2017) Plant-mycorrhizal interactions mediate plant community coexistence by altering resource demand. Ecology 98:187-197. View online.

Mayor J†, Sanders NJ, Classen AT, Bardgett R, Clément JC, Farjado A, Lavorel S, Sundqvist MK, Bahn M, Chisholm C**, Cieraad E, Gedelof Z, Griguilis K, Kudo G, Oberski D, Wardle DA (2017) Elevation alters ecosystem properties across temperate treelines globally. Nature 542:91-95. View online.

Moorhead LC**, Souza L†, Habeck C**, Lindroth RL, Classen AT (2017) Rodent activity slows ecosystem function in an oldfield ecosystem. Ecosphere 8(5):e01777. View online

Souza L†, Stuble KL, Genung MA, Classen AT (2017) Plant genotype identity and intra-specific diversity trump soil nutrient availability to shape old-field structure and function. Functional Ecology 31(4):965-974. View online.

Suonan J**, Classen AT, Zhang Z, He JS (2017) Asymmetric winter warming advanced plant phenology to a greater extent than symmetric warming in an alpine meadow. Functional Ecology 31(11):2147-2156.  View online.

Zhao Q, Classen AT, Wang W, Zhao X, Mao B, Zeng D (2017) Asymmetric effects of litter removal and litter addition on the structure and function of soil microbial communities in a managed pine forest. Plant and Soil 414:81-93. View online.


Classen AT (2016) Editorial. Ecological Monographs. 86:2-3. View online.

Connell RK*, Pfennigwerth AA*, Classen AT, Kwit C (2016) Incorporating redispersal microsites into myrmecochory in eastern North American forests. Ecosphere 7:e01456. View online.

Crowther TW, Todd-Brown KEO, Rowe CW, Wieder WR, Carey JC, Machmuller MB, Snoek LB, Fang S, Zhou G, Allison SD, Blainr JM, Bridgeham SD, Burton AJ, Carrillo Y, Reich PB, Clark JS, Classen AT, Dijkstra FA, Elberling B, Emmett B, Estiarte M, Frey SD, Guo J, Harte J, Jiang L, Johnson BR, Kröel-Dulay G, Larsen KS, Laudon H, Lavallee JM, Luo Y, Lupascu M, Ma LN, Marhan S, Michelsen A, Mohan J, Niu S, Pendell E, Peñuelas J, Pfeifer-Meister L, Poll C, Reinsch S, Reynolds LL, Schmidt IK, Sistla S, Sokol NW, Templer PH, Treseder KK, Welker JM, Bradford MA (2016) Quantifying global soil carbon losses in response to warming. Nature 540:104-108. View online.

Henning, JA**, Weston, DJ, Pelletier, DA, Timm, CM, Jawdy, SS, Classen, AT. (2016) Root bacterial endophytes alter plant phenotype, but not physiology. PeerJ, 4, e2606. View online.

Kuebbing SE**, Patterson C*, Classen AT, Simberloff D (2016) Co-occurring nonnative woody shrubs have additive and non-additive soil legacies. Ecological Applications 26:1896-1906. View online.

Niu S, Classen AT, Dukes JS, Kardol P, Liu L, Lou Y, Rustad L, Vicca S, Tang J, Templer P, Thomas R, Tian D, Wang YP, Xia J, Zaehle S (2016) Global patterns and fundamental mechanisms of the terrestrial nitrogen cycle. Ecology Letters 19:697–709. View online.

Jiang J†, DeAngelis DL, Teh SY, Wang H, Krauss K, Li H, Smith T, Koh HL (2016) Defining the next generation modeling of coastal ecotone dynamics in response to global change. Ecological Modelling  doi:10.1016/j.ecolmodel.2015.04.013. View online.


Classen AT, Sundqvist M†, Henning JA**, Newman GS‡, Moore JAM**, Cregger M†, Moorhead LC**, Patterson CM‡ (2015) ESA Centennial Paper: Direct and indirect effects of climate change on soil microbial and soil microbial-plant interactions: What lies ahead? Ecosphere 6(8):130. View online.

Coble AA, Hart SC, Ketterer ME, Newman GS‡, Kowler AL (2015) Strontium source and depth of uptake shifts with substrate age in semiarid ecosystems. Journal of Geophysical Research 12010691077. DOI: 10.1002/2015JG002992. View online.

De Long JR, Sundqvist MK†, Gundale MJ, Giesler R, Wardle DA (2015) Effects of elevation and nitrogen and phosphorus fertilization on plant defence compounds in subarctic tundra heath vegetation. Functional Ecology, doi: 10.1111/1365-2435.12493. View online.

Ellison AM, Classen AT (2015) Editorial. Ecological Monographs. 85:1-2. View online.

Jing X**, Sanders NJ, Shi Y, Chu H, Classen AT, Zhao K, Chen L, Jiang Y, He JS (2015) The links between ecosystem multifunctionality and above- and belowground biodiversity are mediated by climate. Nature Communications 6:8159. View online.

Kuebbing SE**, Classen AT, Sanders NJ, Simberloff D (2015) The effects of changes in biodiversity differ between phylogenetically paired native and nonnative plant communities. New Phytologist 208(3) 727-735. View online.

Kuebbing SE**, Classen AT, Call J*, Henning J**, Simberloff D (2015) Plant soil interactions promote co-occurrence of three nonnative woody shrubs. Ecology: 96(8): 2289-2299. View online.

Kendrick J*, Ribbons RR**, Classen AT, Ellison AM (2015) Changes in canopy structure and ant assemblages affect soil ecosystem variables as a foundation species declines. Ecosphere 6:art77. View online.

Moore JAM**, Jiang J†, Mayes MA, Wang G, Patterson CM, Classen AT (2015) Interactions among roots, mycorrhizae, and free-living microbial communities differentially impact soil carbon processes. Journal of Ecology 103:1442-1453. View online.

Moore JAM**, Jiang J†, Post WM, Classen AT (2015) Decomposition by ectomycorrhizal fungi alters soil carbon storage in a simulation model. Ecosphere 6(3):29. View online.

Newman GS‡ and Hart SC (2015) Shifting soil resource limitations and ecosystem retrogression across a three million year semi-arid substrate age gradient. Biogeochemistry: 124:177-186. View online.

Siefert A, Violle C, Chalmandrier L, Albert CH, Taudiere A, Fajardo A, Aarssen LW, Baraloto C, Carlucci MB, Cianciaruso MV, Dantas VL, de Bello F, Duarte LDS, Fonseca CR, Freschet GT, Gaucherand S, Gross N, Hikosaka K, Jackson B, Jung V, Kamiyama C, Katabuchi M, Kembel SW, Kichenin E, Kraft NJB, Lagerström A, Bagousse-Pinguet YL, Li Y, Mason N, Messier J, Nakashizuka T, Overton J McC, Peltzer D, Perez-Ramos I M,Pillar VD, Prentice HC, Richardson S, Sasaki T, Schamp BS, Schob C, Shipley B, Sundqvist MK, Sykes MT, Vandewalle M, Wardle DA (2015) A global meta-analysis of the relative extent of intraspecific trait variation in plant communities. Ecology Letters: doi: 10.1111/ele.12508. View online.

Veen GF, Sundqvist MK, Wardle DA (2015) Environmental factors and traits that drive plant litter decomposition do not determine home-field advantage effects. Functional Ecology 29: 981-991. View online.

Veen, GF, Sundqvist, MK (2015) Above-ground and below-ground plant responses to fertilization in two subarctic ecosystem types. Arctic, Antarctic and Alpine Research 47(4), 693-702. View online.


Cregger MA**, McDowell N, Pangle RE, Pockman W, and Classen AT (2014) The impact of precipitation change on nitrogen cycling in a semi-arid ecosystem. Functional Ecology 28: 1534-1544. View online.

Dukes JS, Classen AT, Wan S, Langley JA (2014) Using results from global change experiments to inform land model development and calibration. New Phytologist 204:3. View online.

Schimel D, Strong DR, Ellison AM, Peters DPC, Silver S, Johnson EA, Belnap J, Classen AT, Essington TE, Finley AO, Inouye BD, Stanley EH (2014) Editors are editors, not oracles. Bulletin of the Ecological Society of America 95:342–346. View online.

Fischer DG, Chapman SK, Classen AT, Gehring CA, Grady KC, Schweitzer JA, and Whitham TG (2014) Marschner Review: Plant genetic effects on soils under climate change. Plant and Soil 379: 1-19. View online.

Ellison AM, Classen AT, and Baldwin JD (2014) Editorial. Ecological Monographs. 84:1-2. View online.

Cregger MA**, Sanders NJ, Dunn RR, and Classen AT (2014) Microbial communities respond to experimental warming, but location matters. PeerJ 2:e358. View online.

Barrios-Garcia MN**, Classen AT, and Simberloff D (2014) Disparate responses of above- and belowground properties to soil disturbance by an invasive mammal. Ecosphere 5(4): 44. View online.

Shelton AL, Henning JA**, Schultz P, Clay K (2014) Effects of abundant white-tailed deer on vegetation, animals, mycorrhizal fungi and soils. Forest Ecology and Management 320:39-49. View online.

Wright P*, Cregger MA**, Souza L†, Sanders NJ, and Classen AT (2014) The effects of insects, nutrients, and plant invasion on community structure and function above- and belowground. Ecology & Evolution 4(6):732-742. View online.

Keubbing S**, Classen AT, Simberloff D (2014) Co-occurring invasive woody shrubs alter soil properties and promote exotic species richness. Journal of Applied Ecology 51(1):124-133. View online.

Read QD**, Moorhead LC**, Swenson NJ, Bailey JK, Sanders NJ (2013) Convergent effects of elevation on functional leaf traits within and among species. Functional Ecology 28:37-45. View online.

Sundqvist, M.K., Liu, Z., Giesler, R., Wardle, D.A. (2014) Plant and microbial responses to nitrogen and phosphorus addition across an elevational gradient in subarctic tundra. Ecology 97(7):1819-1835. View online.

Vincent, A.G., Sundqvist, M.K., Wardle, D.A., and Giesler, R. (2014) Bioavailable soil phosphorus decreases with increasing elevation in a subarctic tundra landscape. PLoS ONE 9(3): e92942. View online.

De Long, J.R., Kardol, P., Sundqvist, M.K., Veen, G.F., Wardle, D.A. (2014) Plant growth response to direct and indirect temperature effects varies by vegetation type and elevation in a subarctic tundra. Oikos doi: 10.1111/oik.01764. View online.

Sundqvist, M.K., Wardle, D.A., Vincent, A., Giesler, R. (2014) Contrasting nitrogen and phosphorus dynamics across an elevational gradient for subarctic tundra heath and meadow vegetation. Plant and Soil 383(1):387-399. View online.


Van Nuland ME, Haag EN, Bryant JAM**, Read QD**, Klien RN, Douglas MJ, Gorman CE, Greenwell TD, Busby MW, Collins J, LeRoy JT, Schuchmann G, Schweitzer JA, Bailey JK (2013) Fire promotes pollinator visitations: Implications for ameliorating declines of pollination services. PLoS ONE DOI: 10.1371/journal.pone.0079853. View online.

Burkle LA, Souza L†, Genung MA, Crutsinger GM (2013) Plant genotype, nutrients, and G x E structure floral visitor communities. Ecosphere 4(9):113. View online.

Classen AT, Chapman SK, Whitham TG, Hart SC, and Koch GW (2013) Long-term insect herbivory slows soil development in an arid ecosystem. Ecosphere 4(5):52. View online.

Bryant JAM**, Reynecke BK, and Brown RL. (2013) Geology and topography effects on exotic plant distribution in a semi-arid Mima mound prairie in eastern Washington. Northwest Science 87:12-23. View online.

Kardol P†, Souza L†, Classen AT (2013) Resource availability mediates the importance of priority effects in plant community assembly and ecosystem function. Oikos 122:84-94. View online.


Cregger MA**, Schadt CW, McDowell N, Pockman W, Classen AT (2012) Soil microbial community response to precipitation change in a semi-arid ecosystem. Applied and Environmental Microbiology. 78(24): 8587-8594. View online.

Breza L*, Souza L†, Sanders NJ, and Classen AT (2012) Within and between population variation in plant traits predicts ecosystem functions associated with a dominant plant species. Ecology and Evolution doi: 10.1002/ece3.223. View online.

Hoeksema JD and Classen AT (2012) Is plant genetic control of ectomycorrhizal communities an untapped source of stable soil carbon in managed forests? Plant and Soil 359:197-204. View online.

Castro HF†, Classen AT, Austin EE**, Crawford KM*, Schadt CW (2012) Development and validation of a citrate synthase directed quantitative PCR marker for soil bacterial communities. Applied Soil Ecology 61:69-75. View online.

Del Toro I, Ribbons RR**, Pelini SL (2012) The little things that run the world revisited: a review of ant-mediated ecosystem services and disservices (Hymenoptera: Formicidae). Myrmecological News 17:133-146. View online.

Simberloff D, Souza L†, Nunez MA†, Barrios MN**, Bunn W (2012) The natives are restless, but not often and mostly when disturbed. Ecology 93:598-607. View online.

Lessard JP, Reynolds WN**, Bunn WA, Genung MA, Cregger MA**, Felker-Quinn E, Barrios-Garcia MN, Stevenson ML, Lawton RM, Brown CB, Patrick M, Rock JH, Jenkins MA, Bailey JK, Schweitzer JA (2012) Equivalence in the strength of deer herbivory on above and below ground communities. Basic and Applied Ecology 13:59-66. View online.

Pfennigwerth AA* and Kuebbing SE** (2012) Direct costs associated with invasive non-native plants in Tennessee. Wildland Weeds Summer/Fall: 4-6. View online.


Souza L†, Weston DJ, Sanders NJ, Karve A†, Crutsinger GM†, and Classen AT (2011) Variation from individuals to ecosystems in the response to climatic warming: a test with Solidago altissima. Ecosphere 2(12):1-14. View online.

Gray SB*, Classen AT, Kardol P†, Yermakov Z, and Miller MA (2011) Multiple climate change factors interact to alter soil microbial community structure in an old-field ecosystem. Soil Science Society of America. 75(6): 2217-2226. View online.

Blue JD**, Souza L†, Classen AT, Schweitzer JA, and Sanders NJ (2011) The variable effects of soil nitrogen availability and insect herbivory on aboveground and belowground plant biomass in an old-field ecosystem. Oecologia 167:771-780. View online.

de Graaff MA†, Schadt CW, Rula K*, Six J, Schweitzer JA, and Classen AT (2011) Elevated [CO2] and plant species diversity interact to slow root decomposition. Soil Biology & Biochemistry 43: 2347-2354. View online.

Chen X, Post WM, Norby RJ, and Classen AT (2011) Modeling soil respiration and variation in source components using a multi-factor global climate change experiment. Climatic Change 107(3):459-480. View online.

Classen AT (2011) Book review: Aboveground-belowground linkages: biotic interactions, ecosystem processes, and global change. Richard D. Bardgett and David A. Wardle. Ecology 92(5):1189-1190.

Brosi GB**, McCulley RL, Bush LP, Nelson JA, Classen AT, and Norby RJ (2011) Effects of multiple climate change factors on the tall fescue-fungal endophyte symbiosis: infection frequency and tissue chemistry. New Phytologist 189(3): 797-805. View online.

Garten Jr. CT, Wullschleger SD, and Classen AT (2011) Review and model-based analysis of factors influencing soil carbon sequestration under hybrid poplar. Biomass and Bioenergy 35(1): 214-226. View online.

Iversen CM**, Hooker TD**, Classen AT, and Norby RJ (2011) Net mineralization of N at deeper soil depths as a potential mechanism for sustained forest production under elevated [CO2]. Global Change Biology 17(2):1130-1139. View online.

Kardol P†, Renolds WN**, Norby RJ, and Classen AT (2011) Climate change effects on soil microarthropod abundance and community structure. Applied Soil Ecology 47:37-44. View online.

Luo Y, Melillo JM, Niu S, Beier C, Clark JS, Classen AT, Davidson E, Dukes JS, Evans RD, Field CB, Czimczik CI, Keller M, Kimball BA, Kueppers L, Norby RJ, Pelini SL, Pendall E, Rastetter E, Six J, Smith M, Tjoelker M, Torn M (2011) Coordinated approaches to quantify long-term ecosystem dynamics in response to global change. Global Change Biology 17(2): 843-854. View online.


Castro HF†, Classen AT, Austin EE*, Norby RJ, and Schadt CW (2010) Soil microbial community responses to multiple experimental climate change drivers. Applied and Environmental Microbiology 76(4):999-1007. View online.

de Graaff MA†, Classen AT, Castro H†, and Schadt CW (2010) Labile soil carbon inputs mediate the soil microbial community composition and plant residue decomposition rates. New Phytologist 188:1055-1064. View online.

Classen AT, Norby RJ, Campany CE, Sides KE*, and Weltzin JF (2010) Climate change alters seedling emergence and establishment in an old-field ecosystem. PLoS ONE 5 (10): e13476. View online.

Kardol P†, Cregger MA**, Campany CE, and Classen AT (2010) Soil ecosystem functioning under climate change: plant species and community effects. Ecology 91(3):767-781. View online.

Kardol P†, Campany CE, Souza L†, Norby RJ, Weltzin JF, and Classen AT (2010) Climate change effects on plant biomass alter dominance patterns and community evenness in an experimental old-field ecosystem. Global Change Biology 16: 2676–2687. View online.

Sackett T†, Classen AT, and Sanders NJ (2010) Linking soil food web structure to above- and below-ground ecosystem processes: a meta-analysis. Oikos 119(12):1984-1922. Recommended by Faculty of 1000, 22 Oct 2010. View online.

Matheny PB, Austin EA**, Birkebak JM, Wolfenbarger AD (2010) Craterellus fallax, a Black Trumpet mushroom from eastern North America with a broad host range. Mycorrhiza 20:569-575. View online.


Antoninka A, Wolf JE, Bowker M, Classen AT, and Johnson NC (2009) Linking above and belowground responses to global change at community and ecosystem scales. Global Change Biology 15:914-929. View online.

Austin EE*, Castro HF†, Schadt CW, Sides KT*, and Classen AT (2009) Assessment of 10 years of CO2 fumigation on soil microbial communities and function in a sweetgum plantation. Soil Biology & Biochemistry 41:514-520. View online.

Crutsinger GM**, Sanders NJ, and Classen AT (2009) Comparing intra- and inter-specific effects on litter decomposition in an old-field ecosystem. Basic and Applied Ecology 10:535-543. View online.

Engel EC**, Weltzin JF, Norby RJ, and Classen AT (2009) Responses of an old-field plant community to interacting factors of elevated [CO2], warming, and soil moisture availability. Journal of Plant Ecology 2:1-11. View online.

Garten Jr. CT, Classen AT, and Norby RJ (2009) Watering treatment surpasses elevated CO2 and temperature in importance as a determinant of soil carbon dynamics in a multi-factor climate change experiment. Plant and Soil 319:85-94. View online.


Crutsinger GM**, Habenicht MN*, Classen AT, Schweitzer JA, and Sanders NJ (2008) Galling by Rhopalomyia solidaginis alters Solidago altissima architecture and litter nutrient dynamics in an old-field ecosystem. Plant and Soil 303:95-103. View online.

Crutsinger GM**, Reynolds WN**, Classen AT, and Sanders NJ (2008) Disparate effects of plant genotypic diversity on foliage and litter arthropod communities. Oecologia 158:65-75. View online.

Garten Jr. CT, Classen AT, Norby RJ, Brice DJ, Weltzin JA, and Souza L** (2008) Role of N2-fixation in constructed old-field communities under different regimes of [CO2], temperature, and water availability. Ecosystems 11:125-137. View online.

Hanson PJ, Classen AT, Kueppers L, Luo Y, McDowell NG, Morris J, Rogers A, Thorton P, Ceulemans R, Dukes J, Goulden M, Jackson R, Knapp A, Kirschbaum M, Lewin K, MacCracken M, Melillo J, Ringler T (2008) Ecosystem research: Understanding climate change impacts on ecosystems and feedbacks to the physical climate. Report of the workshop on “Exploring Science Needs for the Next Generation of Climate Change and Elevated CO2 Experiments in Terrestrial Ecosystems” The Department of Energy. View online.

Knapp AK, Beier C, Briske DD, Classen AT, Lou Y, Reichstein M, Smith MD, Smith SD, Bell JE, Fay PA, Heisler JL, Leavitt SW, Sherry R, Smith B, and Weng E (2008) Consequences of more extreme precipitation regimes for terrestrial ecosystems. BioScience 58:811-821. View online.


Classen AT, Chapman SK, Whitham TG, Hart SC, and Koch GW (2007) Genetic-based plant resistance and susceptibility traits to herbivory influence needle and root litter nutrient dynamics. Journal of Ecology 95:1181-1194. View online.

Classen AT, Overby ST, Hart SC, Koch GW, and Whitham TG (2007) Season mediates herbivore effects on litter and soil microbial abundance and activity in a semi-arid woodland. Plant and Soil 295:217-227. View online.

Schadt CW and Classen AT (2007) Book review of: Soil Microbiology, Ecology, and Biochemistry. Eldor A. Paul, editor. Soil Science Society of America 71:1420.

Tyner ML* and Classen AT (2007) Climate change effects on species composition mediates decomposition in an old-field ecosystem. Journal of Undergraduate Research 7:110-114. View online.


Classen AT, DeMarco J*, Hart SC, Whitham TG, Cobb NS, and Koch GW (2006) Impacts of herbivorous insects on decomposer communities during the early stages of primary succession in a semi-arid woodland. Soil Biology & Biochemistry 38:972-982. View online.

Classen AT (2006) Will herbivore range expansions exacerbate the effects of global climate change on soil microclimate? CSA News 51:1. View online.


Classen AT, Hart SC, Whitham TG, Cobb NS, and Koch GW (2005) Insect infestations linked to shifts in microclimate: important climate change implications. Soil Science Society of America Journal 69:2049-2057. View online.

Classen AT and Langley JA (2005) Data-model integration is not magic. New Phytologist 166:367-369. View online.

Hart SC, Classen AT, and Wright RJ (2005) Long-term interval burning alters fine root and mycorrhizal dynamics in a ponderosa pine forest. Journal of Applied Ecology 42:752-761. View online.


Classen AT, Boyle SI, Haskins KE, Overby ST, and Hart SC (2003) Community-level physiological profiles of bacteria and fungi: plate type and incubation temperature influences on contrasting soils. FEMS Microbiological Ecology 44:319-328. View online.

Hart SC and Classen AT (2003) Potential for assessing long-term dynamics in soil nitrogen availability from variations in δ15N of tree rings. Isotopes in Environmental and Health Studies 39:15-28. View online.