Plant and Soil

, Volume 436, Issue 1–2, pp 311–324 | Cite as

Non-additive effects of leaf and twig mixtures from different tree species on experimental litter-bed flammability

  • Weiwei ZhaoEmail author
  • Richard S. P. van Logtestijn
  • Jurgen R. van Hal
  • Ming DongEmail author
  • Johannes H. C. Cornelissen
Regular Article



Tree species can affect litter flammability through leaf size and shape. Larger, simpler-shaped leaf litters form better-ventilated, more flammable litter-beds. However, leaves are generally mixed with twigs in the forest litter layer and together they likely contribute most to surface fire behavior. Here we ask: “Do leaf-twig mixtures have non-additive effects on litter-bed flammability?”


Using laboratory fires, we tested the direction and magnitude of non-additivity of inter- and intra-specific leaf-twig mixtures on litter-bed flammability for four tree species contrasted in leaf size and shape and widespread in fire-prone temperate-boreal forests.


Across species, small needles reduced mixture fuel-bed ignitibility through filling the space between twigs and inhibiting ventilation. Within the small broad-leaved species, the thin, frequently branched and open spaced twigs were too loosely packed to be flammable, while in mixtures the small broad leaves connected these twigs to produce flammable fuel-beds. Once ignited, across species flame spread rate in mixtures was driven by leaves, while fire sustainability was predicted by fuel mass. Fuel-bed flammability was driven more by leaves at larger leaf-to-twig ratio.


For the first time, we demonstrated the existence and mechanisms of non-additive effects of leaf-twig mixtures on experimental litter-bed flammability.


Flammability Leaf Litter mixing Non-additivity Plant traits Surface fire behavior Twig 



We are grateful to the Chinese Scholarship Council for funding WWZ through a 4-year fellowship to study at VU University Amsterdam. The setup of the FLARE laboratory greatly benefitted from grant 047.018.003 by the Netherlands Organization for Scientific Research (NWO) to JHCC. This work was supported by National Key R&D Program of China (grant: 2016YFC0503100) and National Natural Science Foundation of China (grants: 31670429). JHCC and WWZ benefitted from Grant CEP-12CDP007 by the Royal Netherlands Academy of Arts and Sciences (KNAW).

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Supplementary material

11104_2019_3931_MOESM1_ESM.docx (8.3 mb)
ESM 1 (DOCX 8537 kb)


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Copyright information

© Springer Nature Switzerland AG 2019

Authors and Affiliations

  1. 1.Key Laboratory of Hangzhou City for Ecosystem Protection and Restoration, College of Life and Environmental SciencesHangzhou Normal UniversityHangzhouPeople’s Republic of China
  2. 2.Systems Ecology, Department of Ecological Science, Faculty of ScienceVrije Universiteit AmsterdamAmsterdamThe Netherlands

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