Hydraulic traits and tree-ring width in Larix sibirica Ledeb. as affected by summer drought and forest fragmentation in the Mongolian forest steppe

  • Elmira Khansaritoreh
  • Bernhard Schuldt
  • Choimaa Dulamsuren
Original Paper


Key message

Wood-anatomical traits determining the hydraulic architecture of Larix sibirica in the drought-limited Mongolian forest steppe at the southern fringe of the boreal forest respond to summer drought, but only weakly to variations in microclimate that depend on forest stand size.


Siberian larch (L. sibirica Ledeb.) is limited by summer drought and shows increasing mortality rates in the Mongolian forest steppe. The climate sensitivity of stemwood formation increases with decreasing forest stand size. The trees’ hydraulic architecture is crucial for drought resistance and thus the capability to deal with climate warming.


We studied whether hydraulic traits were influenced by temporal or forest size-dependent variations in water availability and were related to tree-ring width.


Hydraulic traits (tracheid diameter, tracheid density, potential sapwood area-specific hydraulic conductivity) of earlywood were studied in stemwood series of 30 years (1985–2014) and were related to climate data. Tree-ring width was measured for the same period. Trees were selected in stands of four different size classes with increasing drought exposure with decreasing stand size.


Tracheid diameters and hydraulic conductivity decreased with decreasing late summer precipitation of the previous year and were positively correlated with tree-ring width. Forest stand size had only weak effects on hydraulic traits, despite known effects on stemwood increment.


Decreasing tracheid diameters and thus hydraulic conductivity are a drought acclimation of L. sibirica in the Mongolian forest steppe. These acclimations occur as a response to drought periods but are little site-dependent with respect to stand size.


Wood anatomy Tracheid diameters Hydraulic conductivity Tracheid density Boreal forest Forest fragmentation 



The study was supported by a grant of the Volkswagen Foundation to M. Hauck, Ch. Dulamsuren and Ch. Leuschner for the project “Forest regeneration and biodiversity at the forest-steppe border of the Altai and Khangai Mountains under contrasting developments of livestock numbers in Kazakhstan and Mongolia”. E. Khansitoreh received an Erasmus Mundus Scholarship in the Salam 2 program. We are thankful to the director of the Tarvagatai Nuruu National Park, Ms. D. Tuya, for her support during field work.


The study was supported by a grant of the Volkswagen Foundation to M. Hauck, Ch. Dulamsuren and Ch. Leuschner for the project “Forest regeneration and biodiversity at the forest-steppe border of the Altai and Khangai Mountains under contrasting developments of livestock numbers in Kazakhstan and Mongolia” (grant no. I/87 175).

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Data availability

The datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request.

Supplementary material

13595_2018_701_MOESM1_ESM.docx (1.3 mb)
ESM 1 (DOCX 1355 kb)


  1. Anfodillo T, Deslauriers A, Menardi R, Tedoldi L, Petit G, Rossi S (2012) Widening of xylem conduits in a conifer tree depends on the longer time of cell expansion downwards along the stem. J Exp Bot 63:837–845. CrossRefPubMedGoogle Scholar
  2. Anfodillo T, Petit G, Crivellaro A (2013) Axial conduit widening in woody species: a still neglected anatomical pattern. IAWA J 34:352–364. CrossRefGoogle Scholar
  3. Begum S, Nakaba S, Oribe Y, Kubo T, Funada R (2010) Changes in the localization and levels of starch and lipids in cambium and phloem during cambial reactivation by artificial heating of main stems of Cryptomeria japonica trees. Ann Bot 106:885–895. CrossRefPubMedPubMedCentralGoogle Scholar
  4. Bryukhanova M, Fonti P (2013) Xylem plasticity allows rapid hydraulic adjustment to annual climatic variability. Trees 27:485–496. CrossRefGoogle Scholar
  5. Buermann W, Parida B, Jung M, MacDonald GM, Tucker CJ, Reichstein M (2014) Recent shift in Eurasian boreal forest greening response may be associated with warmer and drier summers. Geophys Res Lett 41:1995–2002. CrossRefGoogle Scholar
  6. Bunn AG (2008) A dendrochronology program library in R (dplR). Dendrochronologia 26:115–124. CrossRefGoogle Scholar
  7. Carrer M, Von Arx G, Castagneri D, Petit G (2015) Distilling allometric and environmental information from time series of conduit size: the standardization issue and its relationship to tree hydraulic architecture. Tree Physiol 35:27–33. CrossRefPubMedGoogle Scholar
  8. Carter JL, White DA (2009) Plasticity in the Huber value contributes to homeostasis in leaf water relations of a mallee eucalypt with variation to groundwater depth. Tree Physiol 29:1407–1418. CrossRefPubMedGoogle Scholar
  9. Chenlemuge Ts, Dulamsuren Ch, Hertel D, Schuldt B, Leuschner C, Hauck M (2015a) Hydraulic properties and fine root mass of Larix sibirica along forest edge-interior gradients. Acta Oecol 63:28–35CrossRefGoogle Scholar
  10. Chenlemuge Ts, Hertel D, Dulamsuren Ch, Khishigjargal M, Leuschner C, Hauck M (2013) Extremely low fine root biomass in Larix sibirica forests at the southern drought limit of the boreal forest. Flora 208:488–496CrossRefGoogle Scholar
  11. Chenlemuge Ts, Schuldt B, Dulamsuren Ch, Hertel D, Leuschner C, Hauck M (2015b) Stem increment and hydraulic architecture of a boreal conifer (Larix sibirica) under contrasting macroclimates. Trees 29:623–636. CrossRefGoogle Scholar
  12. Cochard H, Froux F, Mayr S, Coutand C (2004) Xylem wall collapse in water-stressed pine needles. Plant Physiol 134:401–408. CrossRefPubMedPubMedCentralGoogle Scholar
  13. Cook E, Holmes R (1984) Program ARSTAN user manual. University of Arizona, Tucson, Arizona, Laboratory of Tree-Ring ResearchGoogle Scholar
  14. Cook E, Kairiukstis L (1990) Methods of dendrochronology. Springer, Dordrecht. CrossRefGoogle Scholar
  15. D’Arrigo R, Jacoby G, Pederson N, Frank D, Buckley B, Nachin B, Mijiddorj R, Dugarjav C (2000) Monogolian tree-rings, temperature sensitivity and reconstructions of Northern Hemisphere temperature. The Holocene 10:669–672. CrossRefGoogle Scholar
  16. De Grandpré L, Tardif JC, Hessl A, Pederson N, Conciatori F, Green TR, Oyunsanaa B, Baatarbileg N (2011) Seasonal shift in the climate responses of Pinus sibirica, Pinus sylvestris, and Larix sibirica trees from semi-arid, north-central Mongolia. Can J For Res 41:1242–1255. CrossRefGoogle Scholar
  17. Delzon S, Douthe C, Sala A, Cohard H (2010) Mechanism of water-stress induced cavitation in conifers: bordered pit structure and function support the hypothesis of seal capillary-seeding. Plant Cell Environ 33:2101–2111. CrossRefPubMedPubMedCentralGoogle Scholar
  18. Domec J-C, Lachenbruch B, Meinzer FC, Woodruff DR, Warren JM, McCulloh KA (2008) Maximum height in a conifer is associated with conflicting requirements for xylem design. Proc Natl Acad Sci U S A 105:12069–12074. CrossRefPubMedPubMedCentralGoogle Scholar
  19. Dulamsuren Ch, Hauck M, Bader M, Osokhjargal D, Oyungerel Sh, Nyambayar S, Runge M, Leuschner C (2009a) Water relations and photosynthetic performance in Larix sibirica growing in the forest-steppe ecotone of northern Mongolia. Tree Physiol 29:99–110CrossRefPubMedGoogle Scholar
  20. Dulamsuren Ch, Hauck M, Bader M, Oyungerel Sh, Osokhjargal D, Nyambayar S, Leuschner C (2009b) The different strategies of Pinus sylvestris and Larix sibirica to deal with summer drought in a northern Mongolian forest-steppe ecotone suggest a future superiority of pine in a warming climate. Can J For Res 39:2520–2528. CrossRefGoogle Scholar
  21. Dulamsuren Ch, Hauck M, Khishigjargal M, Leuschner HH, Leuschner C (2010) Diverging climate trends in Mongolian taiga forests influence growth and regeneration of Larix sibirica. Oecologia 163:1091–1102. CrossRefPubMedPubMedCentralGoogle Scholar
  22. Dulamsuren Ch, Hauck M, Leuschner HH, Leuschner C (2011) Climate response of tree-ring width in Larix sibirica growing in the drought-stressed forest-steppe ecotone of northern Mongolia. Ann For Sci 68:275–282. CrossRefGoogle Scholar
  23. Dulamsuren Ch, Khishigjargal M, Leuschner C, Hauck M (2014) Response of tree-ring width to climate warming and selective logging in larch forests of the Mongolian Altai. J Plant Ecol 7:24–38. CrossRefGoogle Scholar
  24. Dulamsuren Ch, Klinge M, Degener J, Khishigjargal M, Chenlemuge Ts, Bat-Enerel B, Yeruult Yo, Saindovdon D, Ganbaatar Kh, Tsogtbaatar J, Leuschner C, Hauck M (2016) Carbon pool densities and a first estimate of the total carbon pool in the Mongolian forest-steppe. Glob Chang Biol 22:830–844. CrossRefPubMedGoogle Scholar
  25. Dulamsuren Ch, Wommelsdorf T, Zhao F, Xue Y, Zhumadilov BZ, Leuschner C, Hauck M (2013) Increased summer temperatures reduce the growth and regeneration of Larix sibirica in southern boreal forests of eastern Kazakhstan. Ecosystems 16:1536–1549. CrossRefGoogle Scholar
  26. Eckstein D, Bauch J (1969) Beitrag zur Rationalisierung eines dendrochronologischen Verfahrens und zur Analyse seiner Aussagesicherheit. Forstwiss Centralbatt 88:230–250. CrossRefGoogle Scholar
  27. Eilmann B, Weber P, Rigling A, Eckstein D (2006) Growth reactions of Pinus sylvestris L. and Quercus pubescens Willd. to drought years at a xeric site in Valais, Switzerland. Dendrochronologia 23:121–132. CrossRefGoogle Scholar
  28. Eilmann B, Zweifel R, Buchmann N, Fonti P, Rigling A (2009) Drought-induced adaptation of the xylem in Scots pine and pubescent oak. Tree Physiol 29:1011–1020. CrossRefPubMedGoogle Scholar
  29. Fonti P, Babushkina EA (2015) Tracheid anatomical responses to climate in a forest-steppe in southern Siberia. Dendrochronologia 39:32–41CrossRefGoogle Scholar
  30. Fritts HC (1976) Tree rings and climate. Academic Press, LondonGoogle Scholar
  31. Hacke UG, Sperry JS (2001) Functional and ecologycal xylem anatomy. Perspect Plant Ecol Evol Syst 4:97–115. CrossRefGoogle Scholar
  32. Hacke UG, Sperry JS, Pockman WT, Davis SD, McCulloh KA (2001) Trends in wood density and structure are linked to prevention of xylem implosion by negative pressure. Oecologia 126:457–461. CrossRefPubMedGoogle Scholar
  33. Hansen MC, Potapov PV, Moore R, Hancher M, Turubanova SA, Tyukavina A, Thau D, Stehman SV, Goetz SJ, Loveland TR, Kommareddy A, Egorov A, Chini L, Justice CO, Townshend JRG (2013) High-resolution global maps of 21st-century forest cover change. Science 342:850–853. CrossRefPubMedGoogle Scholar
  34. IPCC (2013) Climate change 2013: the physical science basis. Contribution of working group I to the fifth assessment report of the Intergovernmental Panel on Climate Change. Cambridge University Press, CambridgeGoogle Scholar
  35. Jacoby GC, D'Arrigo RD, Davaajamts T (1996) Mongolian tree-rings and 20th-century warming. Science 273:771–773. CrossRefPubMedGoogle Scholar
  36. Khansaritoreh E, Dulamsuren Ch, Klinge M, Ariunbaatar T, Bat-Enerel B, Batsaikhan G, Ganbaatar Kh, Saindovdon D, Yeruult Yo, Tsogtbaatar J, Tuya D, Leuschner C, Hauck M (2017) Higher climate warming sensitivity of Siberian larch in small than large forest islands in the fragmented Mongolian forest steppe. Glob Chang Biol 23:3675–3689. CrossRefPubMedGoogle Scholar
  37. Khishigjargal M, Dulamsuren Ch, Lkhagvadorj D, Leuschner C, Hauck M (2013) Contrasting responses of seedling and sapling densities to livestock density in the Mongolian forest-steppe. Plant Ecol 214:1391–1403. CrossRefGoogle Scholar
  38. Koch GW, Sillett SC, Jennings GM, Davis SD (2004) The limits to tree height. Nature 428:851–854. CrossRefPubMedGoogle Scholar
  39. Liu H, Park Williams A, Allen CD, Guo D, Wu X, Anenkhonov OA, Liang E, Sandanov DV, Yin Y, Qi Z, Badmaeva NK (2013) Rapid warming accelerates tree growth decline in semi-arid forests of Inner Asia. Glob Chang Biol 19:2500–2510. CrossRefPubMedGoogle Scholar
  40. Lkhagvadorj D, Hauck M, Dulamsuren Ch, Tsogtbaatar J (2013) Pastoral nomadism in the forest-steppe of the Mongolian Altai under a changing economy and a warming climate. J Arid Environ 88:82–89. CrossRefGoogle Scholar
  41. Martin-Benito D, Beeckman H, Cañellas I (2013) Influence of drought on tree rings and tracheid features of Pinus nigra and Pinus sylvestris in a mesic Mediterranean forest. Eur J For Res 132:33–45. CrossRefGoogle Scholar
  42. McDowell NG (2011) Mechanisms linking drought, hydraulics, carbon metabolism, and vegetation mortality. Plant Physiol 155:1051–1059. CrossRefPubMedPubMedCentralGoogle Scholar
  43. Oberhuber W, Swidrak I, Pirkebner D, Gruber A (2011) Temporal dynamics of nonstructural carbohydrates and xylem growth in Pinus sylvestris exposed to drought. Can J For Res 41:1590–1597. CrossRefPubMedPubMedCentralGoogle Scholar
  44. Palacio S, Hoch G, Sala A, Körner C, Millard P (2014) Does carbon storage limit tree growth? New Phytol 201:1096–1100. CrossRefPubMedGoogle Scholar
  45. Pennisi E (2005) Tree growth: the sky is not the limit. Science 310:1896–1897. CrossRefPubMedGoogle Scholar
  46. Pittermann J, Sperry JS, Wheeler JK, Hacke UG, Sikkema EH (2006) Mechanical reinforcement of tracheids compromises the hydraulic efficiency of conifer xylem. Plant Cell Environ 29:1618–1628. CrossRefPubMedGoogle Scholar
  47. Poyatos R, Aguadé D, Galiano L, Mencuccini M, Martínez-Vilalta J (2013) Drought-induced defoliation and long periods of near-zero gas exchange play a key role in accentuating metabolic decline of Scots pine. New Phytol 200:388–401. CrossRefPubMedGoogle Scholar
  48. Poyatos R, Martínez-Vilalta J, Čermák J, Ceulemans R, Granier A, Irvine J, Köstner B, Lagergren F, Meiresonne L, Nadezhdina N, Zimmermann R, Llorens P, Mencuccini M (2007) Plasticity in hydraulic architecture of Scots pine across Eurasia. Oecologia 153:245–259. CrossRefPubMedGoogle Scholar
  49. Ryan MG, Phillips N, Bond BJ (2006) The hydraulic limitation hypothesis revisited. Plant Cell Environ 29:367–381. CrossRefPubMedGoogle Scholar
  50. Ryan MG, Yoder BJ (1997) Hydraulic limits to tree height and tree growth: what keeps trees from growing beyond a certain height? Bioscience 47:235–242. CrossRefGoogle Scholar
  51. Schneider L, Gärtner H (2013) The advantage of using a starch based non-Newtonian fluid to prepare micro sections. Dendrochronologia 31:175–178. CrossRefGoogle Scholar
  52. Schweingruber FH, Eckstein D, Serre-Bachet F, Bräker OU (1990) Identification, presentation and interpretation of event years and pointer years in dendrochronology. Dendrochronologia 8:9–38Google Scholar
  53. Sevanto S, Mcdowell NG, Dickman LT, Pangle R, Pockman WT (2014) How do trees die? A test of the hydraulic failure and carbon starvation hypotheses. Plant Cell Environ 37:153–161. CrossRefPubMedGoogle Scholar
  54. Simard S, Giovannelli A, Treydte K, Traversi ML, King GM, Frank D, Fonti P (2013) Intra-annual dynamics of non-structural carbohydrates in the cambium of mature conifer trees reflects radial growth demands. Tree Physiol 33:913–923. CrossRefPubMedGoogle Scholar
  55. Sperry J, Nichols K, Sullivan J, Eastlack S (1994) Xylem embolism in ring-porous, diffuse-porous, and coniferous trees of northern Utah and interior Alaska. Ecology 75:1736–1752. CrossRefGoogle Scholar
  56. Sperry JS, Hacke UG, Pittermann J (2006) Size and function in conifer tracheids and angiosperm vessels. Am J Bot 93:1490–1500. CrossRefPubMedGoogle Scholar
  57. Tei S, Sugimoto A, Yonenobu H, Matsuura Y, Osawa A, Sato H, Fujinuma J, Maximov T (2017) Tree-ring analysis and modeling approaches yield contrary response of circumboreal forest productivity to climate change. Glob Chang Biol 23:5179–5188. CrossRefPubMedGoogle Scholar
  58. Thornthwaite CW (1948) An approach toward a rational classification of climate. Geogr Rev 38:55–94. CrossRefGoogle Scholar
  59. Tsogtbaatar J (2004) Deforestation and reforestation needs in Mongolia. For Ecol Manag 201(1):57–63. CrossRefGoogle Scholar
  60. Tyree MT (2003) Hydraulic limits on tree performance: transpiration, carbon gain and growth of trees. Trees 17:95–100Google Scholar
  61. Tyree MT, Zimmermann MH (2002) Xylem structure and the ascent of sap. Springer, Berlin. CrossRefGoogle Scholar
  62. Vaganov E, Hughes M, Shashkin A (2006) Growth dynamics of conifer tree rings. Springer, BerlinGoogle Scholar
  63. van der Maaten-Theunissen M, van der Maaten E, Bouriaud O (2015) PointRes: an R package to analyze pointer years and components of resilience. Dendrochronologia 35:34–38. CrossRefGoogle Scholar
  64. Vicente-Serrano S, Beguería S, López-Moreno J (2010) A multiscalar drought index sensitive to global warming: the standardized precipitation evapotranspiration index. J Clim 23:1696–1718. CrossRefGoogle Scholar
  65. Wang L, Payette S, Bégin Y (2002) Relationships between anatomical and densitometric characteristics of black spruce and summer temperature at tree line in northern Quebec. Can J For Res 32:477–486. CrossRefGoogle Scholar
  66. White F (1991) Viscous fluid flow. MacGraw, New YorkGoogle Scholar
  67. Wigley TML, Briffa KR, Jones PD, Wigley TML, Briffa KR, Jones PD (1984) On the average value of correlated time series, with applications in dendroclimatology and hydrometeorology. J Clim Appl Meteorol 23:201–213.<0201:OTAVOC>2.0.CO;2 CrossRefGoogle Scholar
  68. Woodruff DR, Bond BJ, Meinzer FC (2004) Does turgor limit growth in tall trees? Plant Cell Environ 27:229–236. CrossRefGoogle Scholar
  69. Yasue K, Funada R, Kobayashi O, Ohtani J (2000) The effects of tracheid dimensions on variations in maximum density of Picea glehnii and relationships to climatic factors. Trees 14:223–229. CrossRefGoogle Scholar
  70. Ziaco E, Biondi F, Rossi S, Deslauriers A (2014) Climatic influences on wood anatomy and tree-ring features of Great Basin conifers at a new mountain observatory. Appl Plant Sci 2, . doi:

Copyright information

© INRA and Springer-Verlag France SAS, part of Springer Nature 2018

Authors and Affiliations

  • Elmira Khansaritoreh
    • 1
  • Bernhard Schuldt
    • 1
  • Choimaa Dulamsuren
    • 1
  1. 1.Plant Ecology, Albrecht von Haller Institute for Plant SciencesGeorg August University of GöttingenGöttingenGermany

Personalised recommendations