Article Accepted Date: 30 August 2014 Acknowledgments This study was supported by the Academy of Finland (grant no. 133495 to Otso Huitu; grants no. 137909 and 110658 to Kari Saikkonen) and by the NERC (grant no. NE/F003994/1 to Xavier Lambin). We thank Dr. Stefan Reidinger and Dr. James Stockdale for help with the silicon analyses. Technician Sinikka Sorsa conducted the phenolic extractions. Stephen Ryan and Anaïs Zimmer assisted with field work.Peer reviewedPublisher PD