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Annual and intra-annual growth dynamics of Myricaria elegans shrubs in arid Himalaya

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To disentangle complex drivers of Myricaria elegans growth in arid Himalaya, we combined tree-ring analysis with detailed dendrometer records. We found that the combination of winter frost, summer floods, and strong summer diurnal temperature fluctuations control annual and intra-annual growth dynamics. The relative importance of these drivers is, however, changing with ongoing climate change.

Abstract

High-mountain areas are among the most sensitive environments to climate change. Understanding how different organisms cope with ongoing climate change is now a major topic in the ecology of cold environments. Here, we investigate climate drivers of the annual and intra-annual growth dynamics of Myricaria elegans, a 3–6 m tall tree/shrub, in a high-elevation cold desert in Ladakh, a rapidly warming region in the NW Himalayas. As Myricaria forms narrow stands around glacier streams surrounded by the desert, we hypothesized that its growth between 3800 and 4100 m will be primarily limited by low temperatures and summer floods. We found that warmer and less snowy conditions in April and May enhance earlywood production. Latewood formation is mostly driven by the June–July temperatures (T). The positive effect of warmer summers on both annual and intra-annual growth is related to fluctuating daily T (from +30 to 0 °C). In particular, dendrometer measurements over a 2-year period showed that net daily growth increments increased when the summer night T remained above 6 °C. While high night T during generally cold desert nights promoted growth, high daytime T caused water stress and growth inhibition. The growth–temperature dependency has gradually weakened due to accelerated warming since the 1990s. In addition, positive latewood responses to high March precipitation during the colder 1960s–1980s have become negative during the warmer 1990s–2000s, reflecting an intensification of summer floods. Latewood width increased while earlywood width decreased from the 1990s, indicating a prolonged growing season and a higher risk of drought-induced embolism in earlywood vessels. Due to a multiplicity of environmental drivers including winter frost, intensified floods and strong summer diurnal T fluctuations, Myricaria growth is not controlled by a single climate parameter. Similar results are increasingly reported from other Himalayan treelines, showing that ongoing climate change will trigger complex and probably spatially variable responses in tree growth. Our study showed that these complex climatic signals can be disentangled by a combination of long-term data from tree-rings with detailed, but short-term, records from dendrometers.

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Acknowledgments

We thank E. Návratová for helping with tree-ring measurements, and Dr. Brian G. McMillian for linguistic help. The study was supported by Czech Science Foundation (GACR 13-13368S), with additional support provided by The Czech Academy of Sciences (long-term research development Project No. RVO 67985939).

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Authors and Affiliations

  1. Institute of Botany, The Czech Academy of Sciences, Zámek 1, 252 43, Průhonice, Czech Republic

    Jiri Dolezal, Eliska Leheckova, Kristina Sohar, Miroslav Dvorsky, Martin Kopecky, Jan Wild & Jan Altman

  2. Department of Botany, Faculty of Science, University of South Bohemia, Na Zlaté stoce 1, 370 05, České Budějovice, Czech Republic

    Jiri Dolezal, Eliska Leheckova & Zuzana Chlumska

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  1. Jiri Dolezal
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Correspondence to Jiri Dolezal.

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Figure A1

Transverse stem section of Myricaria elegans wood made using a sliding microtome and double staining (Astrablue and Safranin) to differentiate lignified (red stained) and unlignified (blue stained) cell walls. Since Myricaria elegans is a ring porous species with an abrupt transition between early and late wood, both parameters are easy to measure and can be used to obtain subseasonal climatic information. Earlywood rings contain one to many rows of pores, more or less compact. Latewood pores are mostly solitary or in small multiples, surrounded by libriform thick-walled fibres. Magnification: 40x, scale bar = 1000 µm (TIFF 17646 kb)

Figure A2

The average temperature (A) and rainfall (B) recorded in the study area for the spring (MAM), summer (JJA), autumn (SON) and winter (DJF) periods, and fitted polynomial regressions (PDF 370 kb)

Figure A3

On the left: Regular cycle lasting approximately 24 hours. Repeating circadian cycles between subsequent local maxima (R_max cycle) and minima (R_min cycle) are distinguished. Differences in radius at starting and ending point of cycles are marked ΔR_max and ΔR_min. On the right: Long cycle lasting more than one day. Steep increase in a stem diameter is followed with several days of a decreasing trend. They are usually caused by rains – plant tissue rehydrates and then slowly desiccates (TIFF 1223 kb)

Figure A4

Comparison of standardized earlywood chronologies of Myricaria elegans between three sites (Rongdu_3800, Rongdu_4100 and Sumur_ 4000) in northern Ladakh, NW Himalayas. A sample depth and a fitted smoothing spline are also shown to depict long-term growth trends (TIFF 1841 kb)

Figure A5

Comparison of standardized latewood chronologies of Myricaria elegans between three sites (Rongdu_3800, Rongdu_4100 and Sumur_ 4000) in northern Ladakh, NW Himalayas. A sample depth and a fitted smoothing spline are also shown to depict long-term growth trends (TIFF 1910 kb)

Figure A6

Comparison of standardized total tree ring width chronologies of Myricaria elegans between three sites (Rongdu_3800, Rongdu_4100 and Sumur_ 4000) in northern Ladakh, NW Himalayas. A sample depth and a fitted smoothing spline are also shown to depict long-term growth trends (TIFF 1852 kb)

Figure A7

Comparison of residual latewood chronologies of Myricaria elegans between three sites (Rongdu_3800, Rongdu_4100 and Sumur_ 4000) in northern Ladakh, NW Himalayas. There was a good agreement of individual minima between the three chronologies (as in the years 2003, 1987, 1976, 1956, 1947) as well as maxima (as in the years 2008, 1982, 1971, 1963, 1943), and significant positive intercorrelations (mean r = 0.44) (TIFF 13118 kb)

Figure A8

Bootsrapped correlation function analysis relating earlywood increments of Myricaria elegans in three sites (Rongdu_3800, Rongdu_4100 and Sumur_ 4000) in NW Himalayas, to monthly mean temperature and precipitation from previous September to current Agust. Abbreviated previous-year months are given in lowercase letters, current year ones in uppercase letters (TIFF 2965 kb)

Figure A9

Bootstrapped correlation function analysis relating latewood width increments of Myricaria elegans in three sites (Rongdu_3800, Rongdu_4100 and Sumur_ 4000) in NW Himalayas, to monthly mean temperature and precipitation from previous September to current Agust. Abbreviated previous-year months are given in lowercase letters, current year ones in uppercase letters (TIFF 3019 kb)

Figure A10

Bootsrapped correlation function analysis relating total ring width increments of Myricaria elegans in three sites (Rongdu_3800, Rongdu_4100 and Sumur_ 4000) in NW Himalayas, to monthly mean temperature and precipitation from previous September to current August. Abbreviated previous-year months are given in lowercase letters, current year ones in uppercase letters (TIFF 2956 kb)

Figure A11

Correlation between the relative change in latewood width in pointer years calculated for the period 1944–2012 and temperature and precipitation in late winter and summer (TIFF 5055 kb)

Figure A12

Average daily stem radius variation and temperature during the vegetation periods in 2010 and 2011. Daily parameters for the first (1.5.–15.6.) and the second (16.6.–31.7.) part of a growth period are displayed on the left and right respectively. Before averaging R_max cycles were detrended by extraction of R_avg_max. Circadian cycles went through a change during the vegetation period. In the first part of the growing season, maximum radius was reached at about 9 a.m. and the trend corresponded with temperature. In the other part of the vegetation season, the maximum set in at about 3 a.m., fluctuations were greater with a more strongly expressed minimum and radius variation possessed an opposite trend to temperature (TIFF 1466 kb)

Table A1

Pointer years at studied localities. △ and ▲ denote positive and negative pointer years respectively (increase or decrease in ring width exceeding 10% of previous ring width were found in at least 70% of trees used in a chronology). △△ and ▲▲ denote strong positive and negative pointer years respectively (relative changes in growth exceeding 15% were present in at least70 % of trees in a chronology) (DOCX 43 kb)

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Dolezal, J., Leheckova, E., Sohar, K. et al. Annual and intra-annual growth dynamics of Myricaria elegans shrubs in arid Himalaya. Trees 30, 761–773 (2016). https://doi.org/10.1007/s00468-015-1318-9

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