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Stronger influence of growth rate than severity of drought stress on mortality of large ponderosa pines during the 2012–2015 California drought

  • Physiological ecology – original research
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Abstract

Forests in the western United States are being subject to more frequent and severe drought events as the climate warms. The 2012–2015 California drought is a recent example, whereby drought stress was exacerbated by a landscape-scale outbreak of western pine beetle (Dendroctonus brevicomis) and resulted in widespread mortality of dominant canopy species including ponderosa pine (Pinus ponderosa). In this study, we compared pairs of large surviving and beetle-killed ponderosa pines following the California drought in the southern Sierra Nevadas to evaluate physiological characteristics related to survival. Inter-annual growth rates and tree-ring stable isotopes (∆13C and δ18O) were utilized to compare severity of drought stress and climate sensitivity in ponderosa pines that survived and those that were killed by western pine beetle. Compared to beetle-killed trees, surviving trees had higher growth rates and grew in plots with lower ponderosa pine basal area. However, there were no detectable differences in tree-ring ∆13C, δ18O, or stable isotope sensitivity to drought-related meteorological variables. These results indicate that differences in severity of drought stress had little influence on local, inter-tree differences in growth rate and survival of large ponderosa pines during this drought event. Many previous studies have shown that large trees are more likely to be attacked and killed by bark beetles compared to small trees. Our results further suggest that among large ponderosa pines, those that were more resistant to drought stress and bark beetle attacks were in the upper echelon of growth rates among trees within a stand and across the landscape.

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References

  • Adams HD, Macalady AK, Breshears DD, Allen CD, Stephenson NL, Saleska SR, Huxman TE, McDowell NG (2010) Climate-induced tree-mortality: earth system consequences. Eos Trans Am Geophysical Union 91:153–154

    Google Scholar 

  • Allen CD, Macalady AK, Chenchouni H, Bachelet D, McDowell N, Vennetier M, Kitzberger T, Riglilng A, Breshears DD, Hogg EH, Gonzalez P, Fensham R, Zhang Z, Castro J, Demidova N, Lim JH, Allard G, Running SW, Semerci A, Cobb N (2010) A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests. For Ecol Manage 259:660–684

    Google Scholar 

  • Amman GD (1972) Mountain pine beetle brood production in relation to thickness of lodgepole pine phloem. J Econ Entomol 65:138–140

    Google Scholar 

  • Anderegg WL, Hicke JA, Fisher RA, Allen CD, Aukema J, Bentz BJ, Hood S, Lichstein JW, Macalady AK, McDowell N, Pan Y, Raffa K, Sala A, Shaw JD, Stephenson NL, Tague C, Zeppel M (2015) Tree mortality from drought, insects, and their interactions in a changing climate. New Phytol 208:674–683

    Google Scholar 

  • Babst F, Bouriaud O, Alexander R, Trouet V, Frank D (2014) Toward consistent measurements of carbon accumulation: A multi-site assessment of biomass and basal area increment across Europe. Dendrochronologia 32(2):153–161

    Google Scholar 

  • Barbour MM, Roden JS, Farquhar GD, Ehleringer JR (2004) Expressing leaf water and cellulose oxygen isotope ratios as enrichment above source water reveals evidence of a Peclet effect. Oecologia 138:426–435

    Google Scholar 

  • Belmecheri S, Wright WE, Szejner P, Morino KA, Monson RK (2018) Carbon and oxygen isotope fractionations in tree rings reveal interactions between cambial phenology and seasonal climate. Plant, Cell Environ 41:2758–2772

    CAS  Google Scholar 

  • Bennett AC, McDowell NG, Allen CD, Anderson-Teixeira KJ (2015) Larger trees suffer most during drought in forests worldwide. Nature Plants 1(10):15139

    Google Scholar 

  • Bentz BJ, Regniere J, Fettig CJ, Hansen M, Hayes JL, Hicke JA, Kelsey RG, Negron JF, Seybold SJ (2010) Climate change and bark beetles of the Western United States and Canada: direct and indirect effects. Science 60:602–613

    Google Scholar 

  • Bracewell RR, Six DL (2014) Broadscale specificity in a bark beetle-fungal symbiosis: a spatio-temporal analysis of the mycangial fungi of the western pine beetle. MicrobEcol 68:859–870

    Google Scholar 

  • Briffa KR, Melvin TM (2011) A closer look at regional curve standardization of tree-ring records: justification of the need, a warning of some pitfalls, and suggested improvements in its application. In: Hughes MK, Diaz HF, Swetnam TW (eds) Dendroclimatology. Springer, Dordrecht, pp 113–145

    Google Scholar 

  • Churchill DJ, Larson AJ, Dahlgreen MC, Franklin JF, Hessburg PF, Lutz JA (2013) Restoring forest resilience: from reference spatial patterns to silvicultural prescriptions and monitoring. For Ecol Manage 291:442–457

    Google Scholar 

  • Cook ER, Krusic PJ (2014) ARSTAN version 44h3: A tree-ring standardization program based on detrending and autoregressive time series modeling, with interactive graphics. Tree-Ring Laboratory, Lamont-Doherty Earth Observatory of Columbia University, Palisades, New York, USA

  • Cook BI, Mankin JS, Marvel K, Williams AP, Smerdon JE, Anchukaitis KJ (2020) Twenty‐first century drought projections in the CMIP6 forcing scenarios. Earth's Future 8(6)

  • Cooper LA, Reed CC, Ballantyne AP (2018) Mountain pine beetle attack faster growing lodgepole pine at low elevations in western Montana, USA. For Ecol Manage 427:200–207

    Google Scholar 

  • Coops NC, Waring RH, Wulder MA, White JC (2009) Prediction and assessment of bark beetle-induced mortality of lodgepole pine using estimates of stand vigor derived from remotely sensed data. Remote Sens Environ 113:1058–1066

    Google Scholar 

  • Craighead FC (1925) Thedendroctonus problem. J Forest 23:340–354

    Google Scholar 

  • Dancho M, Vaughan D (2018) Tidyquant: Tidy quantitative financial analysis. https://bit.ly/2KuBP98

  • Dawson TE, Mambelli S, Plamboeck AH, Templer PH, Tu KP (2002) Stable isotopes in plant ecology. Annu Rev EcolSyst 33:507–559

    Google Scholar 

  • De la Mata R, Hood S, Sala A (2017) Insect outbreak shifts the direction of selection from fast to slow growth rates in the long-lived conifer Pinus ponderosa. ProcNatlAcadSci 114:7391–7396

    Google Scholar 

  • DeMars Jr. CJ, Roettgering BH (1982) Western pine beetle. Forest insect and disease Leaflet 1. USDA forest service, Washington DC, USA

  • Edwards TWD, Fritz P (1986) Assessing meteoric water composition and relative humidity from 18O and 2H in wood cellulose: paleoclimatic implications for southern Ontario. Canada ApplGeochem 1(6):715–723

    CAS  Google Scholar 

  • Ehleringer JR, Hall AE, Farquhar GD (eds) (1993) Stable isotopes and plant carbon-water relations. Academic Press, New York

    Google Scholar 

  • Eldardiry H, Mahmood A, Chen X, Hossain F, Nijssen B, Lettenmaier DP (2019) atmospheric river–induced precipitation and snowpack during the Western United States Cold Season. J Hydrometeorol 20:613–630

    Google Scholar 

  • Farquhar GD, O'Leary MH, Berry JA (1982) On the relationship between carbon isotope discrimination and the intercellular carbon dioxide concentration in leaves. Funct Plant Biol 9(2):121–137

    CAS  Google Scholar 

  • Farquhar GD, Ehleringer JR, Hubick KT (1989) Carbon isotope discrimination and photosynthesis. Annu Rev Plant Physiol Plant MolBiol 40:503–537

    CAS  Google Scholar 

  • Farquhar GD, Lloyd J (1993) Carbon and oxygen isotope effects in the exchange of carbon dioxide between terrestrial plants and the atmosphere. In: Ehleringer JR, Hall AE, Farquhar GD (eds) Stable isotopes and plant carbon-water relations. Academic Press, San Diego, pp 47–70

    Google Scholar 

  • Ferrell RM (2017) Patterns of above-ground ecosystem interactions of regolith in the southern Sierra Nevada of California. Master’s thesis, University of California, Davis, CA, USA

  • Ferrenberg S, Kane JM, Mitton JB (2014) Resin duct characteristics associated with tree resistance to bark beetles across lodgepole and limber pines. Oecologia 174(4):1283–1292

    Google Scholar 

  • Fettig CJ, Mortenson LA, Bulaon BM, Foulk PB (2019) Tree mortality following drought in the central and southern Sierra Nevada, California, U.S. For Ecol Manage 432:164–178

    Google Scholar 

  • Gaylord ML, Kolb TE, Pockman WT, Plaut JA, Yepez EA, Macalady AK, Pangle RE, McDowell NG (2013) Drought predisposes piñon–juniper woodlands to insect attacks and mortality. New Phytol 198:567–578

    CAS  Google Scholar 

  • Gaylord ML, Kolb TE, McDowell NG (2015) Mechanisms of piñon pine mortality after severe drought: a retrospective study of mature trees. Tree Physiol 35(8):806–816. https://doi.org/10.1093/treephys/tpv038

    Article  CAS  Google Scholar 

  • Gleick PH (1987) Regional hydrologic consequences of increases in atmospheric CO2 and other trace gases. Clim Change 10:137–160

    CAS  Google Scholar 

  • Goulden ML, Anderson RG, Bales RC, Kelly AE, Meadows M, Winston GC (2012) Evapotranspiration along an elevation gradient in the Sierra Nevada. J Geophys Res 117:G03028

    Google Scholar 

  • Graf M, Reid ML, Aukema BH, Lindgren BS (2012) Association of tree diameter with body size and lipid content of mountain pine beetles. Can Entomol 144:467–477

    Google Scholar 

  • Griffin D, Anchukaitis KJ (2014) How unusual is the 2012–2014 California drought? Geophys Res Lett 41(24):9017–9023

    Google Scholar 

  • Guerrieri R, Belmecheri S, Ollinger SV, Asbjornsen H, Jennings K, Xiao J, Stocker BD, Martin M, Hollinger DY, Bracho-Garrillo R, Clark K (2019) Disentangling the role of photosynthesis and stomatal conductance on rising forest water-use efficiency. ProcNatlAcadSci 116(34):16909–16914

    CAS  Google Scholar 

  • Helama S, Lindholm M, Timonen M, Eronen M (2004) Detection of climate signal in dendrochronological data analysis: a comparison of tree-ring standardization methods. TheoretApplClimatol 79(3–4):239–254

    Google Scholar 

  • Hessburg PF, Spies TA, Perry DA, Skinner CN, Taylor AH, Brown PM, Stephens SL, Larson AJ, Churchill DJ, Povak NA, Singleton PH (2016) Tamm review: management of mixed-severity fire regime forests in Oregon, Washington, and Northern California. For Ecol Manage 366:221–250

    Google Scholar 

  • Holmes RL (1983) Computer-assisted quality control in tree-ring dating and measurement. Tree Ring Bull 43:69–78

    Google Scholar 

  • Hood S, Sala A (2015) Ponderosa pine resin defenses and growth: metrics matter. Tree Physiol 35:1223–1235

    Google Scholar 

  • Hood S, Sala A, Heyerdahl EK, Boutin M (2015) Low-severity fire increases tree defense against bark beetle attacks. Ecology 96(7):1846–1855

    Google Scholar 

  • Hood SM, Baker S, Sala A (2016) Fortifying the forest: thinning and burning increase resistance to a bark beetle outbreak and promote forest resilience. EcolAppl 0:1–17

  • Jones J (2015) California’s most significant droughts: comparing historical and recent conditions. California Department of Water Resources, Sacramento

    Google Scholar 

  • Kane JM, Kolb TE (2010) Importance of resin ducts in reducing ponderosa pine mortality from bark beetle attack. PhysiolEcol 161:601–609

    Google Scholar 

  • Keen RK (2019) Using tree-ring growth and stable isotopes to explore ponderosa pine ecophysiological responses to climate variability and the 2012–2015 California Drought. Master’s thesis, Utah State University, Logan, UT, USA

  • Kerhoulas LP, Kolb TE, Koch GW (2013) Tree size, stand density, and the source of water used across seasons by ponderosa pine in northern Arizona. For Ecol Manage 289:425–433

    Google Scholar 

  • Kichas NE, Hood SM, Pederson GT, Everett RG, McWethy DB (2020) Whitebark pine (Pinus albicaulis) growth and defense in response to mountain pine beetle outbreaks. For Ecol Manage 457:117736

    Google Scholar 

  • Knapp PA, Soule PT, Maxwell JT (2013) Mountain pine beetle selectivity in old-growth ponderosa pine forests, Montana, USA. EcolEvol 3:1141–1148

    Google Scholar 

  • Kolb TE, Fettig CJ, Ayres MP, Bentz BJ, Hicke JA, Mathiasen R, Steward JE, Weed AS (2016a) Observed and anticipated impacts of drought on forest insects and diseases in the United States. For Ecol Manage 380:321–334

    Google Scholar 

  • Kolb TE, Grady KC, McEttrick MP, Herrero A (2016b) Local-scale drought adaptation of ponderosa pine seedlings at habitat ecotones. For Sci 62(6):641–651

    Google Scholar 

  • Kolb TE, Keefover-Ring K, Burr SJ, Hofstetter R, Gaylord M, Raffa KF (2019) Drought-mediated changes in tree physiological processes weaken tree defenses to bark beetle attack. J ChemEcol 45(10):888–900

    CAS  Google Scholar 

  • Krannitz PG, Duralia TE (2004) Cone and seed production in Pinus ponderosa: a review. Western North America Naturalist 208–218

  • Larson DR, Hann DW (1985) Equations for predicting diameter and squared diameter inside bark at breast height for six major conifers of southwest Oregon. Research Note 77. Forest Research Laboratory, Corvallis, OR, USA

  • Laumer W, Andreu L, Helle G, Schleser GH, Wieloch T, Wissel H (2009) A novel approach for the homogenization of cellulose to use micro-amounts for stable isotope analysis. Rapid Commun Mass Spectrom 23:1934–1940

    CAS  Google Scholar 

  • Leavitt SW, Danzer SR (1993) Method for batch processing small wood samples to holocellulose for stable isotope analysis. Anal Chem 65:87–89

    CAS  Google Scholar 

  • Leavitt SW, Woodhouse CA, Castro CL, Wright WE, Meko DM, Touchan R, Griffin D, Ciancarelli B (2011) The North American monsoon in the US Southwest: potential for investigation with tree-ring carbon isotopes. QuaternInt 235:101–107

    Google Scholar 

  • Margoles DS (2011) Mountain pine beetle-Whitebark pine dynamics in a subalpine ecosystem of the Pioneer mountains, southwest Montana. Master’s thesis. University of Minnesota, Minneapolis, MN, USA

  • McCarroll D, Loader NJ (2004) Stable isotopes in tree rings. QuaternSci Rev 23:771–801

    Google Scholar 

  • McDonald PM (1983) Local volume tables for Pacific madrone, tanoak, and California black oak in north-central California. Research Note PSW-362. USDA Forest Service, Pacific Southwest Forest and Range Experiment Station, Berkeley, CA, USA

  • McDowell M, Brooks JR, Fitzgerald SA, Bond BA (2003) Carbon isotope discrimination and growth response of old Pinus ponderosa trees to stand density reductions. Plant Cell Environ 226:631–644

    Google Scholar 

  • McDowell NG, Adams HD, Bailey JD, Kolb TE (2007) The role of stand density on growth efficiency, leaf area index, and resin flow in southwestern ponderosa pine forests. Can J For Res 37:343–355

    Google Scholar 

  • McDowell NG, Beerling DJ, Breshears DD, Fisher RA, Raffa KF, Stitt M (2011) The interdependence of mechanisms underlying climate-driven vegetation mortality. Trends EcolEvol 26:523–532

    Google Scholar 

  • McLaughlin BC, Ackerly DD, Klos PZ, Natali J, Dawson TE, Thompson SE (2017) Hydrologic refugia, plants, and climate change. Glob Change Biol 23:2941–2961

    Google Scholar 

  • Miller JM, Keen FP (1960) Biology and control of the western pine beetle: a summary of the first fifty years of research. USDA, Washington

    Google Scholar 

  • Moore J, McAfee L, Jirka A, Heath Z, Smith B (2017) 2016 Aerial Survey Results California. Research Paper R5-Pr-034. USDA Forest Service, Pacific Southwest Region, USA

  • Moreira X, Mooney KA, Rasmann S, Petry WK, Carrillo-Gavilán A, Zas R, Sampedro L (2014) Trade-offs between constitutive and induced defenses drive geographical and climatic clines in pine chemical defenses. EcolLett 17:537–546

    Google Scholar 

  • Mote PW, Hamlet AF, Clark MP, Lettenmaier DP (2005) Declining mountain snowpack in western North America. Bull Am Meteor Soc 86:39–50

    Google Scholar 

  • North M, Hurteau M, Fiegener R, Barbour M (2005) Influence of fire and El Nino on tree recruitment by species in Sierran mixed conifer. For Sci 51:187–197

    Google Scholar 

  • Parsons DJ, DeBenedetti SH (1979) Impact of fire suppression on a mixed-conifer forest. For Ecol Manage 2:21–33

    Google Scholar 

  • Person HL (1928) Tree selection by the western pine beetle. J Forest 26:564–578

    Google Scholar 

  • Pile LS, Meyer MD, Rojos R, Roe O, Smith MT (2019) Drought impacts and compounding mortality of forest trees in the southern Sierra Nevada. Forests 10:237

    Google Scholar 

  • Pinheiro J, Bates D, DebRoy S, Sarkar D, R Core Team (2015) nlme: linear and nonlinear mixed effects models. R package version 3.1–121. https://CRAN.R-project.org/package=nlme

  • Raffa KF, Berryman AA (1983) The role of host plant resistance in the colonization behavior and ecology of bark beetles (Coleoptera: Scolytidae). EcolMonogr 53:27–49

    Google Scholar 

  • Raffa KF, Aukema BH, Bentz BJ, Carroll AL, Hicke JA, Turner MG, Romme WH (2008) Cross-scale drivers of natural disturbances prone to anthropogenic amplification: the dynamics of bark beetle eruptions. Bioscience 58:501–517

    Google Scholar 

  • Rigling A, Brühlhart H, Bräker OU, Forster T, Schweingruber FH (2003) Effects of irrigation on diameter growth and vertical resin duct production in Pinus sylvestris L. on dry sites in the central Alps Switzerland. For EcolManag 175(1–3):285–296

    Google Scholar 

  • Rinne KT, Boettger T, Loader NJ, Robertson I, Switsure VR, Waterhouse JS (2005) On the purification of a-cellulose from resinous wood for stable isotope (H, C and O) analysis. ChemGeol 222:75–82

    CAS  Google Scholar 

  • Roden J (2008) Cross-dating of tree ring δ18O and δ13C time series. ChemGeol 252:72–79

    CAS  Google Scholar 

  • Roden JS, Ehleringer LG, JR, (2000) A mechanistic model for interpretation of hydrogen and oxygen isotope ratios in tree-ring cellulose. Geochemica et CosmochimicaActa 64:21–35

    CAS  Google Scholar 

  • Roden JS, Ehleringer JR (2007) Summer precipitation influences the stable oxygen and carbon isotopic composition of tree-ring cellulose in Pinus ponderosa. Tree Physiol 27:491–501

    CAS  Google Scholar 

  • Sevanto S, McDowell NG, Dickman LT, Pangle R, Pockman WT (2014) How do trees die? A test of the hydraulic failure and carbon starvation hypothesis. Plant Cell Environ 37:153–161

    CAS  Google Scholar 

  • Smith GI, Friedman I, Klieforth H, Hardcastle K (1979) Areal distribution of deuterium in eastern California precipitation, 1968–1969. J ApplMeteorol 18:172–188

    CAS  Google Scholar 

  • Stamp N (2003) Out of the quagmire of plant defense hypothesis. Q Rev Biol 78:23–55

    Google Scholar 

  • Stephens SL, Lydersen JM, Collins BM, Fry DL, Meyer MD (2015) Historical and current landscape-scale ponderosa pine and mixed conifer forest structure in the Southern Sierra Nevada. Ecosphere 6(5):1–63

    Google Scholar 

  • Stephenson NL, Das AJ, Ampersee NJ, Bulaon BM, Yee JL (2019) Which trees die during drought? The key role of insect host-tree selection. J Ecol 107(5):2383–2401

    Google Scholar 

  • Stewart IT, Cayan DR, Dettinger MD (2004) Changes in snowmelt runoff timing in western North America under a business as usual climate change scenario. Clim Change 62:217–232

    Google Scholar 

  • Stokes MA, Smiley T (1968) An introduction to tree-ring dating. University of Chicago Press, Chicago

    Google Scholar 

  • Stovall AE, Shugart H, Yang X (2019) Tree height explains mortality risk during an intense drought. Nat Commun 10(1):1–6

    Google Scholar 

  • Szejner P, Wright WE, Babst F, Belmecheri S, Trouet V, Leavitt SW, Ehleringer JR, Monson RK (2016) Latitudinal gradients in tree-ring stable carbon and oxygen isotopes reveal differential climate influences of the North American Monsoon system. J Geophys Res 121:1978–1991

    CAS  Google Scholar 

  • Ulrich DE, Still C, Brooks JR, Kim Y, Meinzer FC (2019) Investigating old-growth ponderosa pine physiology using tree-rings, δ13C, δ18O, and a process-based model. Ecology 100:e02656

    Google Scholar 

  • Van de Water KM, Safford HD (2011) A summary of fire frequency estimates for California vegetation before Euro-American settlement. Fire Ecol 7:26–58

    Google Scholar 

  • Van Mantgem PJ, Stephenson NL, Byrne JC, Daniels LD, Franklin JF, Fule PZ, Harmon ME, Larson AJ, Smith JM, Taylor AH, Veblen TT (2009) Widespread increase of tree mortality rates in the Western United States. Science 323:521–524

    Google Scholar 

  • Voelker SL (2011) Age-dependent changes in environmental influences on tree growth and their implications for forest responses to climate change. In: Meinzer FC, Lachenbruch B, Dawson TE (eds) Size and age-related changes in tree structure and function. Springer, New York, pp 455–479

    Google Scholar 

  • Voelker SL, Meinzer FC, Lachenbruch B, Brooks JR, Guyette RP (2014) Drivers of radial growth and carbon isotope discrimination of bur oak (Quercusmacrocarpa M ichx.) across continental gradients in precipitation, vapour pressure deficit and irradiance. Plant Cell Environ 37(3):766–779

    CAS  Google Scholar 

  • Voelker SL, Merschel AG, Meinzer FC, Ulrich DEM, Spies TA, Still CJ (2019) Fire deficits have increased drought-sensitivity in dry conifer forests: fire frequency and tree-ring carbon isotope evidence from Central Oregon. Glob Change Biol 25:1247–1262

    Google Scholar 

  • Waring RH, Pitman GB (1985) Modifying lodgepole pine stands to change susceptibility to mountain pine beetle attack. Ecology 66:889–897

    Google Scholar 

  • Williams AP, Allen CD, Millar CI, Swetnam TW, Michaelsen J, Still CJ, Leavitt SW (2010) Forest responses to increasing aridity and warmth in the southwestern United States. ProcNatlAcadSci 107(50):21298–21294

    Google Scholar 

  • Williams AP, Allen CD, Macalady AK, Griffin D, Woodhouse CA, Meko DM, Swetnam TW, Rauscher SA, Seager R, Grissino-Mayer HD, Dean JS, Cook ER, Gangodagamage C, Cai M, McDowell NG (2012) Temperature as a potent driver of regional forest drought stress and tree mortality. Nat Clim Change 3:292–297

    Google Scholar 

  • Williams AP, Cook ER, Smerdon JE, Cook BI, Abatzoglou JT, Bolles K, Baek SH, Badger AM, Livneh B (2020) Large contribution from anthropogenic warming to an emerging North American megadrought. Science 368(6488):314–318

    CAS  Google Scholar 

  • Wise EK, Dannenberg MP (2017) Reconstructed storm tracks reveal three centuries of changing moisture delivery to North America. SciAdv 3:e1602263

    Google Scholar 

  • Yanchuk AD, Murphy JF, Wallin KF (2008) Evaluation of genetic variation of attack and resistance in lodgepole pine in the early stages of a mountain pine beetle outbreak. Tree Genet Genomes 4:171–180

    Google Scholar 

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Acknowledgements

We wish to thank the Southern Sierra Critical Zone Observatory (SSCZO) for maintenance of the Soaproot Saddle field site. We specifically thank Erin Stacy from the CZO and Beverly Bulaon from the Forest Service for their assistance with site selection and sampling, and Mike Goulden for substantial information regarding the ecology and history of Soaproot Saddle.

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RMK and SLV designed the experiment and collected data with help from BJB. RMK performed laboratory analyses. RMK and SLV performed data analysis, with substantial input and assistance from SYW and BJB. RF provided additional tree cores from Soaproot Saddle and assisted with data interpretation. All authors significantly contributed to data evaluation and manuscript preparation and gave final approval for publication.

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Correspondence to Rachel M. Keen.

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Communicated by Jim Ehleringer.

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Keen, R.M., Voelker, S.L., Bentz, B.J. et al. Stronger influence of growth rate than severity of drought stress on mortality of large ponderosa pines during the 2012–2015 California drought. Oecologia 194, 359–370 (2020). https://doi.org/10.1007/s00442-020-04771-0

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