, As the simulation results (Figure 1b) suggest, 221 however, consistency in dispersal-model selection for REVEALS and LOVE in the LRA process 222 helps provide robust LRA estimates for reconstruction of the local-to landscape-scale vegetation in 223 spatially and topographically complex terrains, 2015.

, Table 1, Fig. 2) 228 situated between 1600 and 2000 m a.s.l. Seven sites are located on the plateau of Bassiès (Fig. 2) and 229 one site (ARBU) 6 km to the north, We use pollen records from eight small sites: three lakes and five bogs

, All sites are small, varying from 6 to 40 m in radius for bogs and from 60 to 113 m 231 in radius for lakes

. Quintana-seguí, Mean annual temperature in this area (over the 2000-2012 period) is around 7°C and mean 233 annual precipitation about 1640 mm, 2008.

. Szczypta, Winds are mostly coming from the west and north-235 west, November/December and ends in April/May, 2015.

. Theuerkauf, 2017) estimated RPPs for selected 598 taxa in Tasmania. Thus far, the LSM-based estimates of RPPs are available from one study only in 599 NE Germany; considering their variations caused by various environmental and anthropogenic 600 factors, it is necessary to expand the database of the LSM-based RPPs for further applications of the 601 LSM in the LRA reconstruction. All things considered, it is justified for this study to use RPPs for 602 the selected taxa from NW Europe, RPPs need to be obtained from other data sets by 595 assuming the LSM as a pollen dispersal model for consistency, 2012.

, The LRA-based reconstruction of vegetation composition with insect-pollinated plants 606 Plant communities at the mountain tops in our study region are characterized by high 607 abundance of insect-pollinated taxa such as Ericaceae (e.g. Rhododendron spp. and Vaccinium spp.) 608 with semi-dominant grass and Calluna (heather)

. Mazier, 2012); then fall speed of pollen 610 being similar among the constituent taxa, the LRA estimates of Ericaceae, thus the open land cover, 611 tend to increase relative to the other taxa and land cover types. Mazier et al. (2012) excluded 612 zoophilous taxa from the REVEALS reconstruction, mainly because the model assumes that all pollen 613 grains are coming via wind transport. However, the majority of the upland plant taxa common on 614 mountain tops are zoophilous. It is ecologically important to include zoophilous taxa in pollen-based 615 reconstruction of vegetation. Mariani et al. (2017) used both wind-and animal-pollinating plant taxa 616 together for the REVEALS applications in Tasmania successfully, Table 2) relative to the wind-pollinated taxa in general

. Sugita, the LRA result is expressed in proportions of the total cumulative sum of 625 plant abundance at the RSAP (e.g., in biomass, plant cover, foliage mass, basal area, etc.) that is 626 measured in a distance-weighted fashion for all the constituent taxa. Interpretation of the LRA results 627 can be confusing, however. A cumulative sum of the DWPA is a way how pollen records reflect the 628 surrounding vegetation, In theory and practice the RSAP is the smallest spatial scale possible for pollen-based 623 reconstruction of vegetation (Sugita, 1984.

, pollen-vegetation relationship, the site-specific spatial structure of vegetation within the RSAP is 631 impossible to assess quantitatively. Figure 6 is an attempt to visualize how different the spatial 632 structure of the surrounding vegetation at individual sites could be

. Sugita, On the other hand, the 641 LSM predicts higher abundance of long-distance pollen from the regional source, including arboreal 642 plants growing abundantly in lower altitudes, into mountain sites than the GPM does. All these factors, 643 as well as others (e.g., the spatial structure of mountain vegetation, non-random selection of pollen-644 site location, and the lack of RPP estimates considering LSM), The RSAP estimates (Table 3), which are obtained with an inverse modeling of LOVE (Sugita, vol.635, 2007.

, Among the six RSAP estimates listed in Table 3, four estimates are close to or smaller than 2 647 km. The other two, which are obtained with the LSM option, are >4.5 km. Most of the previous 41 648 studies also show that the RSAP estimates are smaller than 2.8 km in radius, when pollen sites are 649 small-to medium-sized, 2010.

. Overballe-petersen, , 2013.

. Poska, In practice, the local 651 vegetation maps are available only within a 2-km radius around each site for comparison, as well, 2014.

, Figure 6 shows that the areas close to the sampling sites are mostly unforested; tree cover 655 (mostly broadleaved trees) increases at and beyond 270-1600 m depending on the sites. The DWPA 656 results depict that

, At the same 659 time it is clear that the spatial structure of vegetation within the RSAP varies among sites significantly, 660 even though the site-to-site variation of the LRA results is small

. Bunting, When the GPM option is selected, the radii of the LRA-based RSAP vary from 663 m (1960-663 1970) to 888 m (1990-2000) to 1038 m, 1994.

, It is reasonable to assume that changes in climate regimes and anthropogenic forcing in the area have 701 was partially supported by a grant from the Estonia Research Council (IUT18-09: the ENCHANTED 702 project)

V. Abrahama, J. Novák, and P. Houfková, A Landscape Reconstruction Algorithm and 708 pedoanthracological data reveal Late Holocene woodland history in the lowlands of the NE Czech 709 Republic, Review of Palaeobotany and Palynology, vol.244, pp.54-64, 2017.

P. G. Appleby, Chronostratigraphic Techniques in Recent Sediments, Last, W.M. & Smol, p.711, 2002.

, Tracking Environmental Change Using Lake Sediments, pp.171-203

D. Allen, A. Simonneau, and G. Le-roux, for publication) Considering lacustrine erosion 714 records and the De Ploey erosion model in an examination of mountain catchment erosion 715 susceptibility and total rainfall reconstruction

M. W. Binford, Calculation and uncertainty analysis of 210Pb dates for PIRLA project lake 717 sediment cores, Journal of Paleolimnology, vol.3, pp.253-267, 1990.

H. H. Birks and H. J. Birks, Future uses of pollen analysis must include plant macrofossils, Journal of Biogeography, vol.45, pp.31-35, 2000.

M. Blaauw, Methods and code for 'classical' age-modelling of radiocarbon sequences, vol.721, 2010.

, Quaternary Geochronology, vol.5, pp.512-518

A. Broström, S. Sugita, and M. J. Gaillard, Pollen productivity estimates for the reconstruction 723 of past vegetation cover in the cultural landscape of southern Sweden. The Holocene, vol.14, pp.368-381, 2004.

A. Broström, S. Sugita, and M. J. Gaillard, Estimating spatial scale of pollen dispersal in the 725 cultural landscape of southern Sweden. The Holocene, vol.15, pp.252-262, 2005.

M. J. Bunting, M. J. Gaillard, and S. Sugita, Vegetation structure and pollen source area. The 727 Holocene, vol.14, pp.651-660, 2004.

A. Claustres, Répartition des éléments traces potentiellement toxiques dans les zones de 729 montagne, Rôle et part des facteurs naturels et anthropiques à l'échelle des temps pédologiques, 2016.

Q. Y. Cui, M. J. Gaillard, and G. Lemdahl, The role of tree composition in Holocene fire 732 history of the hemiboreal and southern boreal zones of southern Sweden, as revealed by the 733 application of the Landscape Reconstruction Algorithm: implications for biodiversity and climate-734 change issues. The Holocene, vol.23, pp.1747-1763, 2013.

Q. Y. Cui, M. J. Gaillard, and G. Lemdahl, Historical land-use and landscape change in 736 southern Sweden and implications for present and future biodiversity, Ecology and Evolution, vol.4, pp.3555-3570, 2014.

F. David, Altitudinal variation in the response of the vegetation to Late-glacial climatic events 739 in the northern French Alps, New Phytologist, vol.125, pp.203-220, 1993.

F. David, Holocene tree limit history in the northern French Alps stomata and pollen evidence. 741 Review of Paleobotany and Palynology, vol.97, pp.227-237, 1997.

D. Macdonald, J. R. Crabtree, and G. Wiesinger, Agricultural abandonment in mountain 743 areas of Europe: Environmental consequences and policy response, Journal of Environmental, vol.744, pp.47-69, 2000.

B. Efron and R. J. Tibshirani, An Introduction to the Bootstrap, 1998.

P. L. Fall, Spatial patterns of atmospheric pollen dispersal in the Colorado Rocky Mountains, 748 USA, Review of Palaeobotany and Palynology, vol.74, pp.293-313, 1992.

R. M. Fyfe, C. Twiddle, and S. Sugita, The Holocene vegetation cover of Britain and Ireland: 750 overcoming problems of scale and discerning patterns of openness, Quaternary Science Reviews, vol.751, pp.132-148, 2013.

M. J. Gaillard, S. Sugita, and M. J. Bunting, The use of modelling and simulation approach 753 in reconstructing past landscapes from fossil pollen data: a review and results from the 754 POLLANDCAL network. Vegetation History and Archaeobotany, vol.17, pp.419-443, 2008.

D. Galop, T. Houet, and F. Mazier, Grazing activities and biodiversity history in the 756 Pyrenees: New insights on high altitude ecosystems in the framework of a Human-Environment 757 Observatory, Past Global Changes Magazine, vol.19, pp.53-55, 2011.

D. Galop, D. Rius, and C. Cugny, Long-term human-environment interactions history in 759 the French Pyrenean Mountains inferred from pollen data, Continuity and 760 Change in Cultural Adaptation to Mountain Environments. Studies in Human Ecology and 761 Adaptation 7, pp.19-30, 2013.

P. A. Herrault, L. Larrieu, and S. Cordier, Combined effects of area, connectivity, history 763 and structural heterogeneity of woodlands on the species richness of hoverflies (Diptera: 764 Syrphidae), Landscape Ecology, vol.31, pp.877-893, 2016.

K. L. Hjelle and S. Sugita, Estimating pollen productivity and relevant source area of pollen 766 using lake sediments in Norway: how does lake size variation affect the estimates? The Holocene, vol.767, pp.313-324, 2012.

K. L. Hjelle, I. K. Mehl, and S. Sugita, From pollen percentage to vegetation cover: 769 evaluation of the Landscape Reconstruction Algorithm in western Norway, Journal of Quaternary, vol.770, pp.312-324, 2015.

K. L. Hjelle, L. S. Halvorsen, and L. Prøsch-danielsen, Long-term changes in regional 772 vegetation cover along the west coast of southern Norway: The importance of human impact, Journal of Vegetation Science, vol.773, pp.404-415, 2018.

T. Houet, L. Vacquié, and F. Vidal, Caractérisation de la fermeture des paysages dans les 775, 2012.

, Pyrénées depuis les années 1940. Application sur le Haut-Vicdessos, vol.33, pp.41-776

T. Houet, L. Vacquié, and D. Sheeren, Evaluating the spatial uncertainty of future land 778 abandonment in a mountain valley, Journal of Mountain Science, vol.12, pp.1-18, 2015.

, Contribution of Working Groups I, II and III 781 to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change, Synthesis Report, 2014.

S. T. Jackson and M. E. Lyford, Pollen dispersal models in Quaternary plant ecology: 784 assumptions, parameters, and prescriptions, The Botanical Review, vol.65, pp.39-75, 1999.

J. Kozak, U. Gimmi, T. Houet, and J. Bolliger, Current practices and challenges for modelling 786 past and future land use and land cover changes in mountainous regions, vol.17, pp.2187-2191, 2017.

A. Kuparinen, T. Markkanen, and H. Riikonen, Modeling air-mediated dispersal of spores, 789 pollen and seeds in forested areas, Ecological Modelling, vol.208, pp.177-188, 2007.

M. Leunda, P. González-sampériz, and G. Gil-romera, The Late-Glacial and Holocene 791, 2017.

, Marboré Lake sequence (2612 m a.s.l., Central Pyrenees, Spain): Testing high altitude sites 792 sensitivity to millennial scale vegetation and climate variability, Global and Planetary Change, vol.793, pp.214-231

M. Leunda, P. González-sampériz, and G. Gil-romera, Ice cave reveals environmental 795 forcing of long-term Pyrenean tree line dynamics, Journal of Ecology, vol.107, pp.814-828, 2019.

M. Mariani, S. Connor, and M. Theuerkauf, Testing quantitative pollen dispersal models in 797 animal-pollinated vegetation mosaics: An example from temperate Tasmania, 2016.

, Quaternary Science Reviews, vol.154, pp.214-225

M. Mariani, P. Kune?, and S. E. Connor, How old is the Tasmanian cultural landscape? A 800 test of landscape openness using quantitative land-cover reconstructions, Journal of Biogeography, vol.801, pp.2410-2420, 2017.

V. Markgraf, Pollen Dispersal in a Mountain Area, Grana, vol.19, pp.127-146, 1980.

L. Marquer, M. J. Gaillard, and S. Sugita, Holocene changes in vegetation composition in 804 northern Europe: why quantitative pollen-based vegetation reconstructions matter, Quaternary 805 Science Reviews, vol.90, pp.199-216, 2014.

L. Marquer, M. J. Gaillard, and S. Sugita, Quantifying the effects of land-use and climate 807 on Holocene plant composition and vegetation change in Europe, Quaternary Science Reviews, vol.808, pp.20-37, 2017.

, Vegetation history. Kluwer, Dordrecht, pp.17-42

P. Quintana-seguí, P. Le-moigne, and Y. Durand, Analysis of Near-Surface Atmospheric 829 Variables: Validation of the SAFRAN Analysis over France, Journal of Applied Meteorology, p.830, 2008.

. Climatology, , vol.47, pp.92-107

P. M. Randall, A study of modern pollen deposition, Review of Palaeobotany and Palynology, vol.64, pp.263-272, 1990.

A. Simonneau, E. Chapron, and T. Courp, Recent climatic and anthropogenic imprints on 834 lacustrine systems in the Pyrenean Mountains inferred from minerogenic and organic clastic 835 supply (Vicdessos valley, vol.23, pp.1762-1775, 2013.

A. Stuart and J. K. Ord, Kendall's advanced theory of statistics, vol.1, 1994.

E. Arnold,

S. Sugita, A model of pollen source area for an entire lake surface, Quaternary Research, vol.39, pp.839-239, 1993.

S. Sugita, Pollen representation of vegetation in Quaternary sediments: Theory and method in 841 patchy vegetation, Journal of Ecology, vol.82, pp.881-897, 1994.

S. Sugita, Theory of quantitative reconstruction of vegetation I: Pollen from large sites 843 REVEALS regional vegetation composition. The Holocene, vol.17, pp.229-241, 2007.

S. Sugita, Theory of quantitative reconstruction of vegetation II: All you need is LOVE. The 845 Holocene, vol.17, pp.243-257, 2007.

S. Sugita, Pollen methods and studies, POLLSCAPE Model: Simulation Approach for Pollen 847 Representation of Vegetation and Land Cover, Encyclopedia of Quaternary 848 Science, pp.871-879, 2013.

S. Sugita, G. M. Macdonald, and C. P. Larsen, Reconstruction of fire disturbance and forest 850 succession from fossil pollen in lake sediments: potential and limitations, p.851, 1997.

, Sediment Records of Biomass Burning and Global Change, pp.852-387

S. Sugita, M. J. Gaillard, and A. Broström, Landscape openness and pollen records: a simulation 854 approach. The Holocene, vol.9, pp.409-421, 1999.

S. Sugita, S. Hicks, and H. Sormunen, Absolute pollen productivity and pollen-vegetation 856 relationships in northern Finland, Journal of Quaternary Science, vol.25, pp.724-736, 2010.

O. G. Sutton, . Micrometeorology, and . Mcgraw-hill, , 1953.

C. Szczypta, S. Gascoin, T. Houet, C. Vigneau, and P. Fanise, Impact of climate and land cover 859 changes on snow cover in a small Pyrenean catchment, Journal of Hydrology, vol.521, pp.84-99, 2015.

M. Theuerkauf, A. Kuparinen, and H. Joosten, Pollen productivity estimates strongly depend 861 on assumed pollen dispersal. The Holocene, vol.23, pp.14-24, 2013.

M. Theuerkauf, J. Couwenberg, and A. Kuparinen, A matter of dispersal: REVEALSinR 863 introduces state-of-the-art dispersal models to quantitative vegetation reconstruction. Vegetation 864 History and Archaeobotany, vol.25, pp.541-553, 2016.

A. K. Trondman, M. J. Gaillard, and F. Mazier, First pollen-based quantitative 866 reconstructions of Holocene regional vegetation cover (plant functional types and land-cover 867 types) in Europe suitable for climate modelling, Global Change Biology, vol.21, pp.676-697, 2015.

A. K. Trondman, M. J. Gaillard, and S. Sugita, Are pollen records from small sites 869 appropriate for REVEALS model-based quantitative reconstructions of past regional vegetation? 870 An empirical test in southern Sweden. Vegetation History and Archaeobotany, vol.25, pp.131-151, 2016.

T. G. Tutin, V. H. Heywood, and N. A. Burgess, -1980) Flora Europaea, 1964.

L. Vacquié, T. Houet, and D. Sheeren, Adapting grazing practices to limit the reforestation 874 of mountainous summer pastures: A process-based approach, Environmental Modelling & 875 Software, vol.84, pp.395-411, 2016.

Y. Zhang, Z. Kong, and Z. Yang, Surface Pollen Distribution from Alpine Vegetation in 877 Eastern Tibet, China. Scientific Reports, vol.7, 2017.

A. Broström, A. B. Nielsen, and M. J. Gaillard, Pollen productivity estimates of key European plant taxa for quantitative reconstruction of past vegetation: a review. Vegetation History and Archaeobotany, vol.17, pp.461-478, 2008.

M. J. Bunting and R. Middleton, Equifinality and uncertainty in the interpretation of pollen data: the Multiple Scenario Approach to reconstruction of past vegetation mosaics. The Holocene, vol.19, pp.799-803, 2009.

M. J. Bunting, M. J. Gaillard, and S. Sugita, Vegetation structure and pollen source area. The Holocene, vol.14, pp.651-660, 2004.

C. Caseldine, R. M. Fyfe, and K. Hjelle, Pollen modelling, palaeoecology and archaeology: virtualization and/or visualisation of the past? Vegetation History and Archaeobotany, vol.17, pp.543-549, 2008.

R. M. Fyfe, GIS and the application of a model of pollen deposition and dispersal: A new approach to testing landscape hypotheses using the POLLANDCAL models, Journal of Archaeological Science, vol.33, pp.483-493, 2006.

R. M. Fyfe, C. Twiddle, and S. Sugita, The Holocene vegetation cover of Britain and Ireland: overcoming problems of scale and discerning patterns of openness, Quaternary Science Reviews, vol.73, pp.132-148, 2013.

M. J. Gaillard, S. Sugita, and M. J. Bunting, The use of modelling and simulation approach in reconstructing past landscapes from fossil pollen data: a review and results from the POLLANDCAL network. Vegetation History and Archaeobotany, vol.17, pp.419-443, 2008.
URL : https://hal.archives-ouvertes.fr/halshs-00961515

S. Hellman, M. J. Gaillard, A. Broström, and S. Sugita, Effects of the sampling design and selection of parameter values on pollen-based quantitative reconstructions of regional vegetation: a case study in southern Sweden using the REVEALS model. Vegetation History and Archaeobotany, vol.17, pp.445-459, 2008.

K. L. Hjelle and S. Sugita, Estimating pollen productivity and relevant source area of pollen using lake sediments in Norway: how does lake size variation affect the estimates? The Holocene, vol.22, pp.313-324, 2012.

A. Kuparinen, T. Markkanen, and H. Riikonen, Modeling air-mediated dispersal of spores, pollen and seeds in forested areas, Ecological Modelling, vol.208, pp.177-188, 2007.

F. Mazier, M. J. Gaillard, and P. Kune?, Testing the effect of site selection and parameter setting on REVEALS-model estimates of plant abundance using the Czech Quaternary Palynological Database, Review of Palaeobotany and Palynology, vol.187, pp.38-49, 2012.
URL : https://hal.archives-ouvertes.fr/halshs-00959845

F. Mazier, A. Broström, and P. Bragée, Two hundred years of land-use change in the South Swedish Uplands: comparison of historical map-based estimates with a pollen-based reconstruction using the landscape reconstruction algorithm, Vegetation History and Archaeobotany, vol.24, pp.555-570, 2015.
URL : https://hal.archives-ouvertes.fr/hal-01171964

I. C. Prentice, Pollen representation, source area, and basin size: toward a unified theory of pollen analysis, Quaternary Research, vol.23, pp.76-86, 1985.

I. C. Prentice, Records of vegetation in time and space: the principles of pollen analysis, Vegetation history, pp.17-42, 1988.

S. Sugita, A model of pollen source area for an entire lake surface, Quaternary Research, vol.39, pp.239-244, 1993.

S. Sugita, Pollen representation of vegetation in Quaternary sediments: Theory and method in patchy vegetation, Journal of Ecology, vol.82, pp.881-897, 1994.

S. Sugita, Theory of quantitative reconstruction of vegetation I: Pollen from large sites REVEALS regional vegetation composition. The Holocene, vol.17, pp.229-241, 2007.

S. Sugita, Theory of quantitative reconstruction of vegetation II: All you need is LOVE. The Holocene, vol.17, pp.243-257, 2007.

S. Sugita, Pollen methods and studies, POLLSCAPE Model: Simulation Approach for Pollen Representation of Vegetation and Land Cover, Encyclopedia of Quaternary Science, pp.871-879, 2013.

S. Sugita, G. M. Macdonald, and C. P. Larsen, Reconstruction of fire disturbance and forest succession from fossil pollen in lake sediments: potential and limitations, Sediment Records of Biomass Burning and Global Change, pp.387-412, 1997.

S. Sugita, M. J. Gaillard, and A. Broström, Landscape openness and pollen records: a simulation approach. The Holocene, vol.9, pp.409-421, 1999.

S. Sugita, S. Hicks, and H. Sormunen, Absolute pollen productivity and pollen-vegetation relationships in northern Finland, Journal of Quaternary Science, vol.25, pp.724-736, 2010.

O. G. Sutton, . Micrometeorology, and . Mcgraw-hill, , 1953.

H. Tauber, Differential pollen dispersion and the interpretation of pollen diagrams, Danmarks Geologiske Undersøgelse. II. RAEKKE, No, vol.89, pp.1-69, 1965.

M. Theuerkauf, A. Kuparinen, and H. Joosten, Pollen productivity estimates strongly depend on assumed pollen dispersal. The Holocene, vol.23, pp.14-24, 2013.

M. Theuerkauf, J. Couwenberg, and A. Kuparinen, A matter of dispersal: REVEALSinR introduces state-of-the-art dispersal models to quantitative vegetation reconstruction. Vegetation History and Archaeobotany, vol.25, pp.541-553, 2016.

A. K. Trondman, M. J. Gaillard, and S. Sugita, Are pollen records from small sites appropriate for REVEALS model-based quantitative reconstructions of past regional vegetation? An empirical test in southern Sweden. Vegetation History and Archaeobotany, REFERENCES Appleby, P.G, vol.25, pp.171-203, 2002.

P. Van-beek, M. Souhaut, and B. Lansard, LAFARA: A new underground laboratory in the French Pyrénées for low-background gamma spectrometry, Journal of Environmental Radioactivity, vol.116, pp.152-158, 2013.

D. Galop, T. Houet, and F. Mazier, Grazing activities and biodiversity history in the Pyrenees: New insights on high altitude ecosystems in the framework of a Human-Environment Observatory, Past Global Changes Magazine, vol.19, pp.53-55, 2011.
URL : https://hal.archives-ouvertes.fr/halshs-00750965

M. E. Goodsite, W. Rom, and J. Heinemeier, High-resolution AMS 14 C dating of post-bomb peat archives of atmospheric pollutants, Radiocarbon, vol.43, pp.495-515, 2001.

S. V. Hansson, A. Claustres, and A. Probst, Atmospheric and terrigenous metal accumulation over 3000 years in a French mountain catchment: Local vs distal influences, Anthropocene, vol.19, pp.45-54, 2017.
URL : https://hal.archives-ouvertes.fr/hal-01760766

D. Mauquoy, M. Blaauw, and B. Van-geel, Late Holocene climatic changes in Tierra del Fuego based on multiproxy analyses of peat deposits, Quaternary Research, vol.61, pp.148-158, 2004.

J. A. Sanchez-cabeza, P. Masqué, and I. Ani-ragolta, Pb and 210 Po analysis in sediments and soils by microwave acid digestion, Journal of Radioanalytical and Nuclear Chemistry, vol.227, pp.19-22, 1998.

A. Simonneau, E. Chapron, and T. Courp, Recent climatic and anthropogenic imprints on lacustrine systems in the Pyrenean Mountains inferred from minerogenic and organic clastic supply (Vicdessos valley, vol.23, pp.1762-1775, 2013.
URL : https://hal.archives-ouvertes.fr/insu-00907274

, Quaternary Science Reviews

. Mainz, , 2019.

, Dear Professor Carrion

, Laurent Marquer and the co-authors

. Dr, ) E-mails: laurent.marquer.es@gmail.com; l.marquer@mpic

L. M. , F. M. , S. S. , and M. , J.G. conceived the initial idea

L. , ran the LRA model

S. S. , led the 3 runs related to the evaluation with simulated landscapes

D. G. , F. M. , and E. ,

S. H. ,

N. D. , F. D. , and G. L. , provided the plant composition in each land-cover type