Skip to main content Accessibility help

Shifts of demography and growth in limber pine forests of the Great Basin, USA, across 4000 yr of climate variability

  • Constance I. Millar (a1), David A. Charlet (a2), Diane L. Delany (a1), John C. King (a3) and Robert D. Westfall (a1)...


Annually dated tree-rings of 509 live and deadwood limber pine (Pinus flexilis) samples from the semi-arid Wassuk Range, Nevada, yielded a 3996-yr record extending from 1983 BC to AD 2013. Correlations of radial growth with climate were positive for water relations and negative for summer temperatures. Long-term trends of ring-width corresponded to climate variability documented from other proxies, including low growth during the Late Holocene Dry Period and Medieval Climate Anomaly (MCA) and elevated growth during cool, wet periods of the Neoglacial and Little Ice Age. Spline fit of the data indicated that growth decrease in the last 20 years was second lowest on record, surpassed by lowest growth at 20 BC—AD 150. Demographics of limber pine by aspect and elevation were not strongly related to long-term climate dynamics, except in the case of extirpations on all but north aspects at the end of the MCA. Pines occurred persistently on north aspects, where a continuous record existed to present. Elevation shifts were not obvious on any aspect, and no evidence existed for migration above current treeline. Non-climatic factors appear to interact with climate to make north slopes refugial for upland pines in semi-arid regions across four millennia.


Corresponding author

*Corresponding author at: Pacific Southwest Research Station, USDA Forest Service, 800 Buchanan St., Albany, California 94710, USA. E-mail address: (C.I. Millar).


Hide All
Adams, K.D., 2007. Late Holocene sedimentary environments and lake-level fluctuations at Walker Lake, Nevada, USA. Geological Society of America Bulletin 119, 126139.
Ashcroft, M.B., 2010. Identifying refugia from climate change. Journal of Biogeography 37, 14071413.
Baker, B.B., Moseley, R.K., 2007. Advancing treeline and retreating glaciers: implications for conservation in Yunnan, PR China. Arctic, Antarctic, and Alpine Research 39, 200209.
Barber, A., 2013. Physiology and Early Life-History Associated with Extreme Longevity: An Investigation of Pinus longaeva (Great Basin bristlecone pine). PhD dissertation, University of California, Santa Cruz.
Beckage, B., Osborne, B., Gavin, D.G., Pucko, C., Siccama, T. Perkins, T., 2008. A rapid upward shift of a forest ecotone during 40 years of warming in the Green Mountains of Vermont. Proceedings of the National Academy of Sciences of the United States of America 105, 41974202.
Bell, D.M., Bradford, J.B., Lauenroth, W.K., 2014. Mountain landscapes offer few opportunities for high‐elevation tree species migration. Global Change Biology 20, 14411451.
Bell, K.L., Johnson, R.E., 1980. Alpine flora of the Wassuk Range, Mineral County, Nevada. Madroño 27, 2535.
Beniston, M., 2003. Climatic change in mountain regions: a review of possible impacts. Climatic Change 59, 531.
Bennett, K.D., Provan, J., 2008. What do we mean by ‘refugia’? Quaternary Science Reviews 27, 24492455.
Benson, L.V., Kashgarian, M., Rye, R.O., Lund, S.P., Paillet, F.L., Smoot, J., Kester, C., Mensing, S., Meko, D., Lindstrom, S., 2002. Holocene multidecadal and multicentennial droughts affecting Northern California and Nevada. Quaternary Science Reviews 21, 659682.
Benson, L.V., Lund, S.P., Smoot, J.P., Kashgarian, M., Burdett, J.W., 2001. Records of climate change from the Owens Lake Basin, California. In: Hershler, R., Madsen, D., Currey, D. (Eds.), Proceedings of the Great Basin Symposium on Glacial and Postglacial Drainage, Smithsonian Institution, Washington, DC.
Bidartondo, M.I., Baar, J., Bruns, T.D., 2001. Low ectomycorrhizal inoculum potential and diversity from soils in and near ancient forests of bristlecone pine (Pinus longaeva). Canadian Journal of Botany 79, 293299.
Birks, H.J., Willis, K.J., 2008. Alpines, trees, and refugia in Europe. Plant Ecology and Diversity 1, 147160.
Bowerman, N.D., Clark, D.H., 2011. Holocene glaciation of the central Sierra Nevada, California. Quaternary Science Reviews 30, 10671085.
Box, G.E.P., Draper, N.R., 1987. Empirical Model-building and Response Surfaces. J. Wiley Sons, New York.
Brubaker, L.B., Anderson, P.M., Edwards, M.E., Lozhkin, A.V., 2005. Beringia as a glacial refugium for boreal trees and shrubs: new perspectives from mapped pollen data. Journal of Biogeography 32, 833848.
Bruns, T.D., Peay, K.G., Boynton, P.J., Grubisha, L.C., Hynson, N.A., Nguyen, N.H., Rosenstock, N.P., 2009. Inoculum potential of Rhizopogon spores increases with time over the first 4 yr of a 99‐yr spore burial experiment. New Phytologist 181, 463470.
Bunn, A.G., 2008. A dendrochronology program library inR (dplR). Dendrochronologia 26, 115124.
Bunn, A.G., Jansma, E., Korpela, M., Westfall, R.D., Baldwin, J., 2013. Using simulations and data to evaluate mean sensitivity (ζ) as a useful statistic in dendrochronology. Dendrochronologia 31, 250254.
Bunn, A.G., Waggoner, L.A., Graumlich, L.J., 2005. Topographic mediation of growth in high elevation foxtail pine (Pinus balfouriana Grev. et Balf.) forests in the Sierra Nevada, USA. Global Ecology and Biogeography 14,103114.
Burns, R.M., Honkala, B.H. (technical coordinators), 1990. Silvics of North America: Volume 1. Conifers. United States Department of Agriculture, Forest Service, Agriculture Handbook 654. Washington, D.C.
Charlet, D.A., 1996. Atlas of Nevada Conifers. A Phytogeographic Reference. University of Nevada Press, Reno.
Collier, F.A., Bidartondo, M.I., 2009. Waiting for fungi: the ectomycorrhizal invasion of lowland heathlands. Journal of Ecology 97, 950963.
Conlisk, E., Castanha, C., Germino, M.J., Veblen, T.T., Smith, J.M., Kueppers, L.M., 2017. Declines in low-elevation subalpine tree populations outpace growth in high-elevation populations with warming. Journal of Ecology 105, 13471357.
Consulting Voortech, 2005. Measure J2X v3.2.1: the tree ring measurement program. Voortech Consulting, Holderness.
Cook, E., Krusic, P., 2014. Program ARSTAN v44h3 (accessed December 15, 2017).
Cook, E.R., Kairukstis, L.A. (Eds.), 1990. Methods of Dendrochronology. Kluwer, Dordrecht.
Coughlan, J.C., Running, S.W., 1997. Regional ecosystem simulation: a general model for simulating snow accumulation and melt in mountainous terrain. Landscape Ecology 12, 119136.
Crimmins, S.M., Dobrowski, S.Z., Greenberg, J.A., Abatzoglou, J.T., Mynsberge, A.R., 2011. Changes in climatic water balance drive downhill shifts in plant species’ optimum elevations. Science 331, 324327.
Daly, C.R., Neilson, R.P., Phillips, D.L., 1994. A statistical-topographic model for mapping climatological precipitation over mountainous terrain. Journal of Applied Meteorology 33, 140158.
Davis, M.B., Shaw, R.G., 2001. Range shifts and adaptive responses to Quaternary climate change. Science 292, 673679.
Dobrowski, S.A., 2011. A climatic basis for microrefugia: the influence of terrain on climate. Global Change Biology 17, 10221035.
Dong, S., Ucarkus, G., Wesnousky, S.G., Maloney, J., Kent, G., Driscoll, N., Baskins, R., 2014. Strike-slip faulting along the Wassuk Range of the northern Walker Lane, Nevada. Geosphere 10, 4048.
Elliott, G.P., 2012. Extrinsic regime shifts drive abrupt changes in regeneration dynamics at upper treeline in the Rocky Mountains, USA. Ecology 93, 16141625.
Esper, J., Cook, E.R., Schweingruber, F.H., 2002. Low-frequency signals in long tree-ring chronologies for reconstructing past temperature variability. Science 295, 22502253.
Felde, V.A., Kapfer, J., Grytnes, J.-A., 2012. Upward shift in elevational plant species ranges in Sikkilsdalen, central Norway. Ecography 35, 922932.
Feng, X., Epstein, S., 1994. Climatic implications of an 8000-year hydrogen isotope time series from bristlecone pine trees. Science 265, 10791081.
Flint, L.E., Flint, A.L., Thorne, J.H., Boynton, R., 2013. Fine-scale hydrologic modeling for regional landscape applications: the California Basin Characterization Model development and performance. Ecological Processes 2, 25.
Fu, P., Rich, P.M., 1999. Design and implementation of the Solar Analyst: an ArcView extension for modeling solar radiation at landscape scales. Proceedings of the Nineteenth Annual ESRI User Conference 1,131.
Gentili, R., Baroni, C., Caccianiga, M., Armiraglio, S., Ghiani, A., Citterio, S., 2015. Potential warm-stage microrefugia for alpine plants: Feedback between geomorphological and biological processes. Ecological Complexity 21, 8799.
Graumlich, L.J., 1993. A 1000-year record of temperature and precipitation in the Sierra Nevada. Quaternary Research 39, 249255.
Grayson, D.K., 2011. The Great Basin. A Natural Prehistory. University of California Press, Berkeley
Griffin, J.R., Critchfield, W.B., 1976. The Distribution of Forest Trees in California. USDA Forest Service Research Paper PSW-82/1972-supplement 1976. US Department of Agriculture, Washington, D.C.
Grissino-Mayer, H.D., 2001. Evaluating crossdating accuracy: a manual and tutorial for the computer program COFECHA. Tree-Ring Research 57, 205221.
Haffer, J., 1982. General aspects of the refuge theory. In: Prance, G.T. (Eds.), Biological Diversification in the Tropics. Columbia University Press, New York, pp. 624.
Hampe, A., Jump, A.S., 2011. Climate relicts: past, present, and future. Annual Review Ecology, Evolution, and Systematics 42, 313333.
Hatchett, B.J., Boyle, D.P., Putnam, A.E. Bassett, S.D., 2015. Placing the 2012–2015 California‐Nevada drought into a paleoclimatic context: Insights from Walker Lake, California‐Nevada, USA. Geophysical Research Letters 42, 86328640.
Holmes, R.L., 1983. Computer-assisted quality control in tree-ring dating and measurement. Tree-Ring Bulletin 43, 6978.
Holmes, R.L., Adams, R.K., Fritts, H.C., 1986. Tree-ring chronologies of western North America: California, Eastern Oregon, and Northern Great Basin with procedures used in the chronology development work including user’s manuals for computer programs COFECHA and ARSTAN. Laboratory of Tree-Ring Research, University of Arizona, Chronology Series VI. Tucson, AZ.
Hughes, M.K., Funkhouser, G., 2003. Frequency-dependent climate signal in upper and lower forest border tree rings in the mountains of the Great Basin. In: Diaz, H. (Ed.) Climate Variability and Change in High Elevation Regions: Past, Present & Future. Springer, Dordrecht, pp. 233244.
Huntley, B., Birks, J.H., 1983. An Atlas of Past and Present Pollen Maps for Europe 0–13,000 Years Ago. Cambridge University Press, Cambridge.
Huntley, B., Webb, T. III, 1989. Migration: species’ response to climatic variations caused by changes in the earth’s orbit. Journal of Biogeography 16, 519.
International Tree-Ring Data Bank (ITRDB), 2017–2018. NOAA National Climate Data Center (accessed February 19, 2018).
Jeffress, M.R., VanGunst, J., Millar, C.I., 2017. A surprising discovery of American pika sites in the northwest Great Basin. Western North American Naturalist 77, 252268.
Konrad, S.K., Clark, D.H., 1998. Evidence for an early Neoglacial glacier advance from rock glaciers and lake sediments in the Sierra Nevada, California, U.S.A. Arctic and Alpine Research 30, 272284.
Krusic, P., Cook, E., 2013. ARSTAN40 for Mac OS, v44h2 (accessed August 8, 2017).
LaMarche, V. Jr., 1973. Holocene climatic variations inferred from treeline fluctuations in the White Mountains, California. Quaternary Research 3, 632660.
LaMarche, V. Jr., 1974. Paleoclimatic inferences from long tree-ring records. Science 183, 10431048.
LaMarche, V. Jr., Mooney, H.A., 1967. Altithermal treeline advance in western United States. Nature 213, 980982.
Legendre, P., Legendre, L., 1998. Numerical Ecology. 2nd English ed. Elsevier, New York.
Lenoir, J., Ge’gout, J.C., Guisan, A., Vittoz, P., Wohlgemuth, T., Zimmerman, N.E., Dullinger, S., Pauli, H., Willner, W., Svenning, J.C., 2010. Going against the flow: potential mechanisms for unexpected downslope range shifts in a warming climate. Ecography 33, 295303.
Lenoir, J., Ge’gout, J.C., Marquet, P.A., Ruffray, P., Brisse, H., 2008. A significant upward shift in plant species optimum elevation during the 20th century. Science 320, 17681771.
Lloyd, A.H., Graumlich, L.J., 1997. Holocene dynamics of treeline forests in the Sierra Nevada. Ecology 78, 11991210.
Lorenz, T.J., Sullivan, K.A., Bakian, A.V., Aubry, C.A., 2011. Cache-site selection in Clark’s Nutcracker (Nucifraga columbiana). The Auk 128, 237247.
Louderback, L.A., Rhode, D.E., 2009. 15,000 years of vegetation change in the Bonneville basin: the Blue Lake pollen record. Quaternary Science Review 28, 308326.
Mensing, S., Benson, L.V., Kashgarian, M., Lund, S., 2004. A Holocene pollen record of persistent droughts from Pyramid Lake, Nevada, USA. Quaternary Research 62, 2938.
Mensing, S.A., Sharpe, S.E., Tunno, I., Sada, D.W., Thomas, J.M., Starratt, S., Smith, J., 2013. The Late Holocene Dry Period: multiproxy evidence for an extended drought between 2800 and 1850 cal yr BP across the central Great Basin, USA. Quaternary Science Reviews 78, 266282.
Mensing, S.A., Smith, J., Norman, K.B., Allan, M., 2008. Extended drought in the Great Basin western North America in the last two millennia reconstructed from pollen records. Quaternary International 188, 7989.
Millar, C.I., Charlet, D.A., Westfall, R.D., King, J.C., Delany, D.L., Flint, A., Flint, L.E., 2018. Do low-elevation ravines provide climate refugia for subalpine limber pine (Pinus flexilis) in the Great Basin, USA? Canadian Journal of Forest Research 48, 663671.
Millar, C.I., King, J.C., Westfall, R.D., Alden, H.A., Delany, D.L., 2006. Late Holocene forest dynamics, volcanism, and climate change at Whitewing Mountain and San Joaquin Ridge, Mono County, Sierra Nevada, CA, USA. Quaternary Research 66, 273287.
Millar, C.I., Westfall, R.D., Delany, D.L., 2007. Response of high-elevation limber pine (Pinus flexilis) to multiyear droughts and 20th-century warming, Sierra Nevada, California, USA. Canadian Journal of Forest Research 37, 25082520.
Millar, C.I., Westfall, R.D., Delany, D.L., Flint, A.L., Flint, L.E., 2015. Recruitment patterns and growth of high-elevation pines in response to climatic variability (1883–2013), in the western Great Basin, USA. Canadian Journal of Forest Research 45, 12991312.
Millar, C.I., Westfall, R.D., Delany, D.L., King, J.C., Graumlich, L.C., 2004. Response of subalpine conifers in the Sierra Nevada, California, U.S.A. to 20th-century warming and decadal climate variability. Arctic, Antarctic, and Alpine Research 36, 181200.
Miller, J., House, K., Germanoski, D., Tausch, R., Chambers, J., 2004. Fluvial geomorphic responses to Holocene climate change. In: Chambers, J.C., Miller, J.R. (Eds.), Great Basin Riparian Ecosystems: Ecology, Management and Restoration. Island Press, Covelo, pp. 4987.
Morelli, T.L., Daly, C., Dobrowski, S.Z., Dulen, D.M., Ebersole, J.L., Jackson, S.T., Lundquist, J.D., et al., 2016. Managing climate change refugia for climate adaptation. PLoS One 11,
Moritz, C., Patton, J.L., Conroy, C.J., Parra, J.L., White, G.C., Beissinger, S.R., 2008. Impact of a century of climate change on small-mammal communities in Yosemite National Park, USA. Science 322, 261264.
Osborne, G., Bevis, K., 2001. Glaciation in the Great Basin of the western United States. Quaternary Science Reviews 20, 13771410.
R Core Team, 2017. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
Rapacciuolo, G., Maher, S.P., Schneider, A.D., Hammond, T.T., Jabis, M., Walsh, R.E., Iknayan, K.J., et al., 2014. Beyond a warming fingerprint: individualistic biogeographic responses to heterogeneous climate change in California. Global Change Biology 20, 28412855.
Reinemann, S.A., Porinchu, S.F., Bloom, A.M., Mark, B.G., Box, J.E., 2009. A multiproxy paleolimnological reconstruction of Holocene climate conditions in the Great Basin, United States. Quaternary Research 72, 347358.
Rull, V., 2009. Microrefugia. Journal of Biogeography 36, 481484.
Salzer, M.W., Bunn, A.G., Graham, N.E., Hughes, M.K., 2014. Five millennia of paleotemperature from tree-rings in the Great Basin, USA. Climate Dynamics 42, 15171526.
Salzer, M.W., Hughes, M.K., Bunn, A.G., Kipfmueller, K.F., 2009. Recent unprecedented tree-ring growth in bristlecone pine at the highest elevations and possible causes. Proceedings National Academy of Science of the United States of America 106, 2034820353.
SAS Institute, 2015. SAS Online, Version 12. JMP Statistics and Graphics Guide, Cary.
Scuderi, L.A., 1993. A 2000-year tree ring record of annual temperatures in the Sierra Nevada Mountains. Science 259, 14331436.
Serra-Diaz, J.M., Scheller, R.M., Syphard, A.D., Franklin, J., 2015. Disturbance and climate microrefugia mediate tree range shifts during climate change. Landscape Ecology 30, 10391053.
Smith, S.E., Read, D.J., 2008. Mycorrhizal Symbiosis. 3rd ed. Academic Press, London.
Steele, R.G.D., Torrie, J.H., 1980. Principles and Procedures of Statistics. McGraw Hill, New York.
Stine, S., 1990. Late Holocene fluctuations of Mono Lake, eastern California. Paleogeography, Palaeoclimatology, Palaeoecology 78, 333381.
Stine, S., 1994. Extreme and persistent drought in California and Patagonia during mediaeval time. Nature 369, 546549.
Strachan, S., Daly, C., 2017. Testing the daily PRISM air temperature model on semiarid mountain slopes. Journal of Geophysical Research: Atmospheres 122, 56975715.
Surpless, B.E., 2012. Cenozoic tectonic evolution of the central Wassuk Range, western Nevada, USA. International Geology Review 54, 547571.
Tausch, R., Nowak, C., Mensing, S., 2004. Climate change and associated vegetation dynamics during the Holocene: the paleoecological record. In: Chambers, J.C., Miller, J.R. (Eds.), Great Basin Riparian Ecosystems: Ecology, Management and Restoration. Island Press, Covelo, pp. 2448.
Thompson, R.S., 1992. Late Quaternary environments in Ruby Valley, Nevada. Quaternary Research 37, 115.
Toy, T.J., Foster, G.R., Renard, K.G., 2002. Soil Erosion: Processes, Prediction, Measurement, and Control. John Wiley and Sons, Hoboken, N.J.
Van Gunst, K.J., Weisberg, P.J., Yang, J., Fan, Y., 2016. Do denser forests have greater risk of tree mortality: a remote sensing analysis of density-dependent forest mortality. Forest Ecology and Management 359, 1932.
Wolfram Research, 2017. Mathematica, Version 11.1. Wolfram Research, Inc., Champaign.


Type Description Title
Supplementary materials

Millar et al. supplementary material
Figure S1

 Unknown (955 KB)
955 KB
Supplementary materials

Millar et al. supplementary material
Figure S2

 Unknown (522 KB)
522 KB
Supplementary materials

Millar et al. supplementary material
Figure S3

 Unknown (1.0 MB)
1.0 MB
Supplementary materials

Millar et al. supplementary material
Millar et al. supplementary material 1

 Word (14 KB)
14 KB

Shifts of demography and growth in limber pine forests of the Great Basin, USA, across 4000 yr of climate variability

  • Constance I. Millar (a1), David A. Charlet (a2), Diane L. Delany (a1), John C. King (a3) and Robert D. Westfall (a1)...


Altmetric attention score

Full text views

Total number of HTML views: 0
Total number of PDF views: 0 *
Loading metrics...

Abstract views

Total abstract views: 0 *
Loading metrics...

* Views captured on Cambridge Core between <date>. This data will be updated every 24 hours.

Usage data cannot currently be displayed.