[1] Univ Helsinki, Dept Forest Sci, POB 27, FIN-00014 Helsinki, Finland
[2] Finnish Geospatial Inst, Ctr Excellence Laser Scanning Res, Geodeetinrinne 2, Masala 02431, Finland
[3] Finnish Geospatial Inst, Dept Remote Sensing & Photogrammetry, Geodeetinrinne 2, Masala 02431, Finland
[4] Univ Eastern Finland, Sch Forest Sci, POB 111, Joensuu 80101, Finland