Supplementary Materials

1. Broadleaved-evergreen biotype

LandClim calculates the biomass of species cohorts, and uses separate allometric relationships to convert the biomass of coniferous and broadleaved-deciduous species to stem diameter (Schumacher et al. 2004). We added a new biomass to diameter conversion for broadleaved-evergreen species based on scaling functions for mature trees and the wood density of Quercus ilex (Pilli et al. 2006), the dominant broadleaved-evergreen species in Italian forests:

(1)

Where a = 0.12817 and is a scaling coefficient based on the density of Q. ilex (0.95 t m-3) and b = 2.5 and is a scaling exponent for mature trees, which is consistent with our means for calculating growth rates and maximum size.

LandClim tracks light availability using leaf area index. Each species is assigned one of five foliage types based on crown density that relate diameter to foliage biomass (Supplemental Table 2; Bugmann 1994; Bugmann 1996). We assumed that the ratio of dry to wet foliage mass, and the specific leaf area of broadleaved-evergreen species are the mean of coniferous and broadleaved-deciduous species (Fyllas et al. 2007).

2. Estimation of life history parameters

LandClim simulates vegetation dynamics on the basis of species-specific life history parameters (Table 2). Plant growth is based on a maximum potential growth rate (rs, tons yr-1) under optimal conditions for each species. Growth is reduced from this maximum by growth reduction functions determined by light availability, drought stress, and growing degree days (Schumacher et al. 2004). We estimated the maximum potential growth rate (rs) using yield tables from Italian forests (Castellani 1982; Federici et al. 2001) to rank the growth rates of commercially-relevant Mediterranean species (e.g. Quercus cerris, Q. ilex) relative to species that were previously parameterized for LandClim. For non-commercial species, we estimated growth rates based on physiological studies from the Mediterranean region (e.g. Baldocchi et al. 2010; Ogaya and Penuelas 2007; Paraskevopoulos et al. 1994; Sanchez-Gomez et al. 2008). We determined maximum potential biomass from height and diameter measurements from a database of Italian big trees (data source: Associazione Ricerche Documentazioni Etologiche e Ambientali, Rome, Italy) following methodology described by Schumacher and Bugmann (2006).

Seedling establishment and growth in LandClim are limited by winter cold and summer warmth using the mean temperature of the coldest month, and the mean annual total of growing degree days, respectively. We estimated temperature constraints for the new Mediterranean species by overlaying spatially-interpolated climatic data from the period 1961-1990 (New et al. 2001) with species distribution maps (EUFORGEN 2009a, b; Meusel et al. 1964; Scholzel et al. 2002), and assigned the minima as two standard deviations below the mean value for the range of each species (Table 2). For species without digitized distribution maps, we generalized temperature tolerances from bioindicator values (Pignatti 2005). We estimated the sensitivity to fire damage to trees based on measurements of stem survival following fire (Fernandes et al. 2008; Gonzalez et al. 2007), and assumed all shrubs have low tolerance to fire. We determined resprouting capacity from a database of fire-related traits of plant species of the Mediterranean Basin (Paula et al. 2009).

Previous LandClim implementations for Central Europe ranked the shade and drought tolerance of each species on a scale of 1 (highly intolerant) – 5 (highly tolerant; Henne et al. 2011; Schumacher and Bugmann 2006). Because the Mediterranean flora includes species that are more adapted to drought stress than those in Central Europe, and because woody species typical of open sunny environments are common in the Mediterranean, we extended both scales to 1 – 6 using bioindicator values (Ellenberg 1996; Pignatti 2005). This strategy provides continuity in the environmental tolerance parameters between Central Europe and the Mediterranean.

3. Estimating the probability of fire spread and browsing mortality

We used the incomplete beta function to relate the probability of fire spread to the LandClim drought index, and the probability of establishment to browsing pressure. The incomplete beta function is defined by the integral (Press et al. 1992):

(2) where α and β are > 0,

(3) and B(α, β) is the value of the beta function:

The parameters α and β define the shape of the function and x is the normalized value of the drought index or browsing intensity.

For all variations in the fire regime, β was constant (i.e. 1.5). For simulations with small infrequent fires, α = 2.5. We increased the probability of fire spread in simulations with large infrequent fires, and small frequent fires by setting α = 2.0. Because fires also occur during dry intervals of wet years, especially when humans increase the number of ignition sources, we set the minimum probability to 0.1 for fire regimes with small infrequent, and large infrequent fires, and to 0.17 for the small frequent fire regime. We used a maximum fire probability of 0.8 for all fire regimes to avoid unrealistically-high fire spread values.

We simulated vegetation dynamics with three browsing intensities: without browsing (x = 0), low browsing (x = 0.25), and moderate browsing (x = 0.5). Each species is assigned a browsing tolerance value (i.e. brTol, Table 2) on a scale of 1 (very intolerant to browsing) to 5 (very tolerant to browsing). A separate cumulative beta probability density function is defined for each browsing tolerance: brTol 1, α = 4, β = 12; brTol 2, α = 2.4, β = 3.5; brTol 3, α = 1.9, β = 1.6; brTol 4, α = 2, β = 1; brTol 5, α = 4, β = 1.


Supplemental Table 1. List of trees and shrubs included in simulations.

Abies alba

Acer campestre

Acer monspessulanum

Acer platanoides

Acer pseudoplatanus

Arbutus unedo

Carpinus betulus

Castanea sativa

Cistus salvifolius

Corylus avellana

Erica arborea

Fagus sylvatica

Fraxinus excelsior

Fraxinus ornus

Ilex aquifolium

Laurus nobilis

Olea europea

Ostrya carpinifolia

Phillyrea angustifolia

Phillyrea latifolia

Pinus halepensis

Pistacia lentiscus

Pistacia terebinthus

Quercus cerris

Quercus coccifera

Quercus ilex

Quercus petraea

Quercus pubescens

Quercus robur

Quercus suber

Rhamnus alaternus

Tilia cordata

Tilia platyphyllos

Ulmus glabra


Supplemental Table 2a. Life history parameters for Mediterranean species not included in previous LandClim versions, and other important species mentioned in text.

Species r K maxAge matu ED MD vegP spAge

(t yr-1) (t) (yr) (yr) (m) (m) (yr)

Abies alba 0.08 17.6 700 70 50 160 0 0

Arbutus unedo 0.07 0.7 400 20 30 -1 1 75

Cistus salvifolius 0.005 0.005 70 10 40 170 0 0

Erica arborea 0.06 0.1 100 10 30 100 1 50

Fraxinus ornus 0.07 2.7 200 20 40 140 1 50

Laurus nobilis 0.06 0.5 300 20 30 -1 1 50

Olea europea 0.04 10.0 700 10 30 -1 1 75

Ostrya carpinifolia 0.1 2.0 150 30 55 180 1 30

Phillyrea latifolia 0.03 0.2 300 10 30 -1 1 50

Phillyrea angustifolia 0.03 0.2 50 10 30 -1 1 50

Pinus halepensis 0.07 2.7 275 20 90 300 0 0

Pistacia lentiscus 0.04 0.4 300 10 30 -1 1 50

Pistacia terebinthus 0.04 0.4 300 10 30 -1 1 50

Quercus cerris 0.09 11.6 600 60 30 -1 1 75

Quercus coccifera 0.03 1.4 450 20 30 -1 1 75

Quercus ilex 0.03 8.9 600 20 30 -1 1 75

Quercus pubescens 0.07 2.4 500 60 30 -1 1 50

Quercus suber 0.03 8.3 400 20 30 -1 1 75

Rhamnus alaternus 0.005 0.01 100 10 30 -1 1 50

r: maximum above-ground biomass growth rate; K: maximum above-ground tree biomass; maxAge: maximum longevity; matu: age for seed production; ED: effective seeding distance; MD: maximum seeding distance; vegP: species capable (1) or not capable (0) of resprouting following disturbance; spAge: maximum age for vegetative reproduction. Data sources are listed in the text.

Table 2b.

Species BioT folType shTol minDD mint drTol brTol FireTol

(d) (°C)

Abies alba NE 5 6 641 -6 3 1 3

Arbutus unedo BE 4 1 1792 2 5 3 1

Cistus salvifolius BE 4 1 1792 2 6 4 1

Erica arborea BE 4 3 1554 2 5 5 1

Fraxinus ornus D 2 4 1639 0 5 3 2

Laurus nobilis BE 5 6 1800 0 1 3 1

Olea europea BE 4 1 2031 6 6 3 3

Ostrya carpinifolia D 2 5 937 -7 4 4 2

Phillyrea latifolia BE 4 4 2074 1 4 5 1

Phillyrea angustifolia BE 4 1 2174 2 6 5 1

Pinus halepensis NE 4 1 2303 2 6 3 3

Pistacia lentiscus BE 4 1 2031 2 6 4 1

Pistacia terebinthus D 4 1 1554 0 6 4 1

Quercus cerris D 2 3 1554 -1 4 2 3

Quercus coccifera BE 4 3 1963 1 5 4 3

Quercus ilex BE 4 6 1773 1 5 3 3

Quercus pubescens D 3 3 1011 -99 5 2 3

Quercus suber BE 4 5 2635 6 5 3 5

Rhamnus alaternus BE 4 5 1792 2 6 3 1

BioT: NE denotes needle-leaved evergreen, BE broad-leaved evergreen, and D deciduous species. folType: Foliage type; shTol: shade tolerance (1 = very intolerant, 6 = very tolerant). To expand the range for the Mediterranean species, we added 1 to the values used by Schumacher (2004), but the implementation is the same in the model for like species. minDD: minimum annual degree-day sum; mint: minimum temperature for establishment; drTol: drought tolerance (1 = very intolerant, 6 = very tolerant); brTol: browsing tolerance (1 = very tolerant, 5 = very intolerant); FireTol: sensitivity to fire damage of trees that have reached the canopy.


References

Baldocchi D.D., Ma S.Y., Rambal S., Misson L., Ourcival J.M., Limousin J.M., Pereira J. and Papale D. 2010. On the differential advantages of evergreenness and deciduousness in mediterranean oak woodlands: a flux perspective. Ecological Applications 20: 1583-1597.

Bugmann H. 1994. On the ecology of mountainous forests in a changing climate: a simulation study, ETH, Zurich, Switzerland. 258pp.

Bugmann H.K.M. 1996. A simplified forest model to study species composition along climate gradients. Ecology 77: 2055-2074.

Castellani C. 1982. Tavole Stereometriche ed Alsometriche Costuite per Boschi Italiani. Istituto Sperimentale per l'Assestamento Forestale e per l'Alpicoltura, Trento.

Ellenberg H. 1996. Vegetation Mitteleuropas mit den Alpen in ökologischer Sicht. E. Ulmer, Stuttgart.

EUFORGEN 2009a. Distribution map of Aleppo pine (Pinus halepensis). Available online at www.euforgen.org.

EUFORGEN 2009b. Distribution map of cork oak (Quercus suber). Available online at www.euforgen.org.

Federici S., Quaratino R., Papale D., S. T. and Valentini R. 2001. Informatic system of Italy yield tables. DiSAFRi - University of Tuscia.

Fernandes P.M., Vega J.A., Jiménez E. and Rigolot E. 2008. Fire resistance of European pines. Forest Ecology and Management 256: 246-255.

Fyllas N.M., Phillips O.L., Kunin W.E., Matsinos Y.G. and Troumbis A.I. 2007. Development and parameterization of a general forest gap dynamics simulator for the North-eastern Mediterranean Basin (GREek FOrest Species). Ecological Modelling: 439-456.

Gonzalez J.R., Trasobares A., Palahi M. and Pukkala T. 2007. Predicting stand damage and tree survival in burned forests in Catalonia (North-East Spain). Annals of Forest Science 64: 733-742.

Henne P.D., Elkin C.M., Reineking B., Bugmann H. and Tinner W. 2011. Did soil development limit spruce (Picea abies) expansion in the Central Alps during the Holocene? Testing a palaeobotanical hypothesis with a dynamic landscape model. Journal of Biogeography 38: 933-949.

Meusel H., Jäger E., Rauschert S. and Weinert E. 1964. Vergleichende Chorologie der zentraleuropäischen Flora Band 1, Karten. Fischer, Jena.

New M., Todd M., Hulme M. and Jones P. 2001. Precipitation measurements and trends in the twentieth century. International Journal of Climatology 21: 1899-+.

Ogaya R. and Penuelas J. 2007. Tree growth, mortality, and above-ground biomass accumulation in a holm oak forest under a five-year experimental field drought. Plant Ecology 189: 291-299.

Paraskevopoulos S.P., Iatrou G.D. and Pantis J.D. 1994. Plant-growth strategies in evergreen-sclerophyllous shrublands (maquis) in central Greece. Vegetatio 115: 109-114.

Paula S., Arianoutsou M., Kazanis D., et al. 2009. Fire-related traits for plant species of the Mediterranean Basin. Ecology 90: 1420-1420.

Pignatti S. 2005. Valori di bioindicazione delle piante vascolari della flora d'Italia. Braun-Blanquetia 39: 1-97.

Pilli R., Anfodillo T. and Carrer M. 2006. Towards a functional and simplified allometry for estimating forest biomass. Forest Ecology and Management 237: 583-593.

Press W.H., Teukolsky S.A., Vetterling W.T. and Flannery B.P. 1992. Numerical Recipes in C: the art of scientific computing. Cambridge University Press.

Sanchez-Gomez D., Zavala M.A., Van Schalkwijk D.B., Urbieta I.R. and Valladares F. 2008. Rank reversals in tree growth along tree size, competition and climatic gradients for four forest canopy dominant species in Central Spain. Annals of Forest Science 65: Article No.: 605.

Scholzel C.A., Hense A., Hubl P., Kuhl N. and Litt T. 2002. Digitization and geo-referencing of botanical distribution maps. Journal of Biogeography 29: 851-856.

Schumacher S. 2004. The role of large-scale disturbances and climate for the dynamics of forested landscapes in the European Alps, Swiss Federal Institute of Technology: Zürich, Switzerland.

Schumacher S. and Bugmann H. 2006. The relative importance of climatic effects, wildfires and management for future forest landscape dynamics in the Swiss Alps. Global Change Biology 12: 1435-1450.

Schumacher S., Bugmann H. and Mladenoff D.J. 2004. Improving the formulation of tree growth and succession in a spatially explicit landscape model. Ecological Modelling 180: 175-194.

Schumacher S., Reineking B., Sibold J. and Bugmann H. 2006. Modeling the impact of climate and vegetation on fire regimes in mountain landscapes. Landscape Ecology 21: 539-554.