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The authors confirm that all the data that underlie the results presented in this study are available within the Supplementary Information files and can be downloaded in conjunction with this paper.

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The data were funded and provided by the CNRS TRAM Project, ANR-15-CE01-0008, Observatoire Homme-Milieu Pyrénées Haut Vicdessos-LABEX DRIIHM ANR-11-LABX0010 and CESBIO. The research leading to these results has also received funding from the People Programme (Marie Curie Actions) of the European Union’s Seventh Framework Programme (FP7/2007-2013) under REA grant agreement no. PCOFUND-GA-2013-609102, through the PRESTIGE programme coordinated by Campus France. The authors acknowledge that this work was carried out in the CMAC National Facility, housed within the University of Strathclyde’s Technology and Innovation Centre, who are funded with a UKRPIF (UK Research Partnership Institute Fund) capital award, SFC ref. H13054, from the Higher Education Funding Council for England (HEFCE).

Author information

These authors contributed equally: S. Allen, D. Allen.


EcoLab (Laboratoire Ecologie Fonctionnelle et Environnement), ENSAT, UMR-CNRS 5245, Castanet Tolosan, France
Steve Allen
, Deonie Allen
, Gaël Le Roux
, Pilar Durántez Jiménez
& Stéphane Binet

Department of Civil and Environmental Engineering, University of Strathclyde, Glasgow, UK
Steve Allen
& Vernon R. Phoenix

ISTO, CNRS UMR 7327, Université d’Orléans, BRGM, Orléans, France
Anaëlle Simonneau
& Stéphane Binet

GEODE, UMR-CNRS 5602, Université Toulouse Jean Jaurès, Toulouse, France

S.A. and D.A. designed the study, undertook all the analyses and co-authored the manuscript. G.L.R. and V.R.P. provided the study design and analytical guidance and assisted in the preparation and revision of the manuscript. P.D. undertook all the field sampling and field protocol design, assisted in the sample preparation and contributed to the manuscript. A.S., S.B. and D.G. provided financial support and field site access that enabled this study to occur and contributed to the manuscript.

Competing interests

The authors declare no competing interests.

Corresponding author

Correspondence to Deonie Allen.

Supplementary information

Supplementary Description, Supplementary Figures 1-3 and Supplementary Table 1

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