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<p> ESM_Changes in range limits under climate change Page 1 of 12</p><p>Supplementary Material</p><p>Study systems</p><p>We investigated how climate change and population dynamics together affect the range </p><p> limits of two lagomorphs of conservation concern: the Mexican Volcano Rabbit </p><p>5(Romerolagus diazi) and the European Mountain Hare (Lepus timidus). Romerolagus </p><p> diazi is protected by national and international legislation: listed as an endangered species</p><p> in the Mexican law (DOF 2001) and in the IUCN Red List (LSG 1996). Lepus timidus is </p><p> listed under Annex V of the EC Habitats Directive (1992), which implies that a number </p><p> of methods of capture are restricted or banned (JNCC 2007). </p><p>10</p><p>The current distribution of Romerolagus diazi is at its range limit (see Fig. S1 </p><p> supplementary material), due to historical non-climatic drivers of range contraction. </p><p>Although the species has always been endemic to a small region of the Transvolcanic </p><p>Belt in central Mexico, between 2,800 and 4,250 meters above sea level (Romero & </p><p>15Velázquez 1999; Velázquez et al. 1999) it was previously more widespread. A total </p><p> occupancy area of about 280 km2 has been estimated in a discontinuous distribution </p><p> pattern, from which four major core areas have been identified (Velázquez et al. 1996). </p><p>Population size at the end of the 1980’s was estimated at between 2,478 and 12,120 </p><p>(Velázquez 1994). Besides its natural restricted distribution, several factors threaten the </p><p>20long-term persistence of Romerolagus diazi. First, two of its core areas are located about </p><p>30 km south of Mexico City -one of the largest urban areas in the world- imposing a </p><p> strong pressure to the rabbit’s habitat, which has been quickly reducing in the last </p><p> decades due to urbanization and expansion of the grazing and cropland frontier, the ESM_Changes in range limits under climate change Page 2 of 12</p><p> increase of frequency of wildfires, illegal hunting, and introduction of domestic dogs and </p><p>25cats that prey upon the rabbits are among the most important ones (Hoth et al. 1989; </p><p>Romero & Velázquez 1999). Second, more recent climatic changes have been detected in</p><p> the distributional area of Romerolagus diazi, in particular an increase in the winter </p><p> temperature of more than 1.5 °C in the last 40 years (Domínguez 2007). It is this </p><p> combination of pressures that makes this species highly vulnerable to extinction.</p><p>30</p><p>Fig. S1. Distribution of Romerolagus diazi (blue) superimposed over the 2010 climate </p><p> suitability.</p><p>Lepus timidus is an arctic/subartic species with a fragmented range across Europe, </p><p>35extending across Russia in the east and to Scotland and Ireland in the west (Mitchell-</p><p>Jones et al. 1999; Thulin 2003). In Europe the northwest range limit of this species is in ESM_Changes in range limits under climate change Page 3 of 12</p><p>Scotland. It is highly fragmented and although Lepus timidus is capable of existing in a </p><p> wide range of habitats and environmental conditions, in Great Britain (England, Wales </p><p> and Scotland) it is generally associated with heather (Calluna and Erica spp., Newey et al.</p><p>402007). The current distribution of Lepus timidus populations show generally regular but </p><p> sometimes dramatic changes in density, depending on habitat suitability related to </p><p> vegetation and climate [see below] (Newey et al. 2007). The current distribution of Lepus</p><p> timidus within Britain, the geographical area of interest in the current paper, is estimated at</p><p>76 721 km2 (JNCC 2007). </p><p>45</p><p>Habitat suitability</p><p>Romerolagus diazi inhabits coniferous forests with a dense sub-alpine grassland cover in </p><p> the herbaceous stratum. It is highly sensitive to vegetation structure and composition, </p><p> thus perturbations to structural habitat has an important impact in its populations </p><p>50(Cervantes et al. 1990; Cervantes & Martínez 1996). The habitat map for the Volcano </p><p>Rabbit was obtained from the last land use / land cover map generated for the whole </p><p> country in 2000 (IGUNAM-INEGI 2001). Original land cover classes were reclassified in</p><p> highly suitable (1), suitable (0.5), and unsuitable (0) for the Volcano Rabbit based on </p><p>Velázquez et al. (1999).</p><p>55</p><p>Habitat for Lepus timidus was based on the proportion of suitable habitat in the cell using </p><p> the 1990 Land Cover Map (Fuller et al. 1994) sub categories that are largely heather, </p><p> open shrub moor (10) and shrub heath (25). In the absence of more detailed information </p><p> on the relationship between hare density and vegetation cover, we here assumed that grid ESM_Changes in range limits under climate change Page 4 of 12</p><p>60cells were potentially suitable habitat if the percentage cover of heather exceeded 70%. </p><p>After applying this vegetation mask (categorical threshold), the bioclimatic was thereafter</p><p> used to determine relative habitat suitability.</p><p>Occurrence data </p><p>Occurrence data for the Volcano Rabbit was obtained from four scientific collections </p><p>65[Colección Nacional de Mamíferos, Instituto de Biología, UNAM (CNMA); Museum of </p><p>Natural History, University of Kansas (KU); Instituto Politécnico Nacional (IPN) and </p><p>Colección de Mamíferos of Universidad Autónoma Metropolitana (Unidad Iztapalapa) </p><p>(UAMI)], scientific literature (Velázquez et al. 1996), and from fieldwork carried out </p><p> from 2004-2007 {Domínguez, 2007 #1262}.</p><p>70</p><p>Presences for the Mountain Hare (Lepus timidus) were based on records held at the </p><p>Biological Records Centre (BRC) for Great Britain (www.nbn.org.uk/). Squares (10km </p><p> x 10km) within Great Britain with no record at or below 10km resolution were </p><p> considered an absence. For simplicity, all 1km squares within a 10km square were </p><p>75considered a potential occupancy, provided the total habitat suitability threshold </p><p>(vegetation + climate – see below) was >0.2.</p><p>Historic distribution of Romerolagus diazi</p><p>Occurrence data for the Romerolagus diazi were obtained from four scientific collections </p><p>80(see above). Historical distribution map was generated using species distribution </p><p> modelling techniques and GIS analysis. A potential distribution map of the species was </p><p> created using a maximum entropy approach implemented in the MaxEnt software 5ESM_Changes in range limits under climate change Page 5 of 12</p><p>(Phillips et al. 2006). For the purpose of this study, we used all available occurrences of </p><p> the species and a suite of 19 bioclimatic variables produced specifically for Mexico with </p><p>85the ANUSPLIN software (Hutchinson 1997) and kindly provided by O. Tellez-Valdes. </p><p>Automatic settings were used in Maxent for all tuning parameters following Phillips and </p><p>Dudik (2008) and we selected the new logistic output format to obtain a map that is </p><p> interpreted as an estimate of the probability of presence. The probabilistic map was </p><p> converted into a binary map (presence/absent) selecting as threshold the minimum </p><p>90probability value in which all occurrences were predicted. Finally, those climatically </p><p> suitable areas which have never been inhabited by the species due to dispersal limitation </p><p> or historical reasons were trimmed based on distributional range limits described in </p><p>Cervantes et al. (1990) and Velázquez et al. (1996).</p><p>Climate suitability methods </p><p>95Generalized Additive Models (GAMs) with binomial errors, a logit link and a smoothed </p><p> function (3 nodes) were used to build the bioclimatic models (mgcv (1.4-1) library in R, </p><p>Wood 2008). GAMs are semi-parametric models with data-driven response curves </p><p>(Hastie 1992). We took an ensemble modelling approach using a multi-model inference </p><p> framework (Burnham & Anderson 2002; Araujo & New 2007; Thuiller et al. 2007) based</p><p>100on all-subsets selection of the GAMs using the AIC measure. The ten ‘best’ models </p><p>(lowest AIC) were conserved and the final projections were a weighted average of these.</p><p>Romerolagus diazi</p><p>The Mexican volcano rabbit model was built and projected on a 0.01° x 0.01° (~1km2) </p><p> grid. This is a very range-restricted species and the number of presences was small </p><p>105compared to the number of absences. Rather than using all the absences in the ESM_Changes in range limits under climate change Page 6 of 12</p><p> surrounding area (>180 000 points) we chose a random set of absences. Bioclimate </p><p> modelling was the same as for the mountain hare except that due to the smaller number of</p><p> presences only three bioclimatic variables were used to build the models; Annual Mean </p><p>Temperature, Temperature Seasonality (standard deviation *100), Annual Precipitation. </p><p>110These three variables were highly correlated (r >0.6) with a larger set of variables </p><p> including those used in the mountain hare modelling. All 7 possible models (i.e., one </p><p> model for each unique combination of the three bioclimate variables) were used and were</p><p> weighted as in the mountain hare modelling (see below).</p><p>Lepus timidus</p><p>115The mountain hare (Lepus timidus) bioclimatic model was built on the ~50 x 50km Atlas </p><p>Flora Europea (AFE) grid, using presence absence distribution data from the European </p><p>Mammals Atlas (Mitchell-Jones et al. 1999) as the response variable. We used four </p><p> bioclimatic variables that have been previously been shown to correspond well to species </p><p> range limits: mean temperature of the warmest month and coldest month, annual </p><p>120precipitation, and the ratio of actual to potential evapotranspiration (Thuiller et al. 2005; </p><p>Levinsky I. et al. 2007). Climate data used to build the model was based on the 10’ </p><p>European grid for the last recognized climate normal period (1961-1990) generated by </p><p> the FP5 ATEAM project (New et al. 2002; Schroter et al. 2005). These data were </p><p> aggregated to 50 x 50 km Universal Transverse Mercator (UTM) in ArcGIS/ArcInfo 9.2 </p><p>125(ESRI, Redlands, California, USA) to match the species data grid. The model was then </p><p> projected onto the 10 x 10km British National Grid (BNG) using climate data for the </p><p> same variables for both the recent past and future (1961-2100). ESM_Changes in range limits under climate change Page 7 of 12</p><p>Future projections</p><p>Future projections for the climate variables were derived using climate model outputs </p><p>130from the HadCM3 global climate model made available through the Intergovernmental </p><p>Panel on Climate Change (IPCC) Data Distribution Centre (ipcc-ddc.cru.uea.ac.uk). </p><p>Modelled climate anomalies were scaled based on the A2 storyline (Special Report on </p><p>Emissions Scenarios: http://www.grida.no/climate/ipcc/emission/); this scenario describes</p><p> a heterogeneous future world focused on self-reliance, preservation of local identities and</p><p>135slower and more regional economic development (Nakicenovic & Swart 2000). This </p><p> scenario is more conservative than the A1FI scenario, which we are currently tracking </p><p> well above (Rahmstorf et al. 2007), but A2 better reflects emissions scenarios now </p><p> gaining favour among mitigation policy researchers, encompassing a later shortage of </p><p> fossil fuels and more active mitigation policies which come into force by mid century </p><p>140(www.ipcc.ch). Given the likelihood of currently unmodelled slow feedbacks in the </p><p> climate system (Hansen et al. 2007), it is plausible that future climate change will be </p><p> more extreme than that used here, so we consider our simulations to be a mid-range </p><p> projection.</p><p>145Future projections for Romerolagus diazi were calculated as an averaged projection </p><p> surface for each of three future 30year means provided by the IPCC (2010-2039, 2040-</p><p>2069, and 2070-2099). For interpolation purposes these were labelled for the first year </p><p> and the interpolation extended 30 years to 2100. For the dynamic climate maps these </p><p> values were then interpolated to annual time slices using (“gi_beta_r44.exe” Ersts & R. ESM_Changes in range limits under climate change Page 8 of 12</p><p>1502008). Grid interpolator applies a linear interpolation between two grids on a cell by cell </p><p> basis. </p><p>InterpolationIncrementi,j = EndYeari,j - StartYeari,j / ( EndYeari,j - StartYeari,j )</p><p>For each year between the StartYear and EndYear</p><p>CurrentYeari,j = StartYeari,j + ( InterpolationIncrementi,j * ( CurrentYear - StartYear ).</p><p>155</p><p>Future climate envelope projections for Lepus timidus were produced in the same way </p><p>(using 30-year means 1991-2020, 2021-50, 2051-80). These were then converted to a </p><p>1 x 1 km grid to match the habitat and occurrence layer for Great Britain, but with all </p><p> squares within the HadCM3-modelled 10 km having the same climate value. The annual </p><p>160interpolation was extended by 49 years to 2100.</p><p>Stochastic Population Models</p><p>Romerolagus diazi and Lepus timidus population models comprised 3 stage classes - </p><p> leveret; one year old; and those two years or greater. Demographic rates (survival, age of </p><p>165maturity and fecundity rates) were based directly on published estimates or taken from </p><p> similar species (Flux & Angerman 1990; Hewson & Hinge 1990; Marboutin et al. 2003; </p><p>Newey et al. 2004; Dahl 2005; Jennings et al. 2006; Mahony & Montgomery 2006). </p><p>Leveret survival was estimated at 30-50% and adult survival at 45-70%. Age of maturity </p><p> was modelled at 1 year (Iason 1989) and fecundity was related to age, with an average </p><p>170litter size of 5 for 2-year olds and 6 for 3-year olds. We estimated variability in vital rates </p><p> by trialling values until the fluctuations in the simulated populations matched those of the</p><p> time series. ESM_Changes in range limits under climate change Page 9 of 12</p><p>Density Dependence</p><p>Density dependence (DD) was implemented using a Scramble model (Logistic or Ricker </p><p>175type of DD), which determined the population growth rate at each time step (by </p><p> modifying fecundity and leveret survival) as a function of the population size at that time </p><p> step. Maximum rate of population growth (Rmax) was calculated by fitting exponential </p><p> and logistic models to time series of abundances from the Global Population Dynamics </p><p>Database (http://www3.imperial.ac.uk/cpb/research/patternsandprocesses/gpdd). AIC </p><p>180model averaging was used to provide a weighted estimate of maximum/intrinsic </p><p> population growth rate for 17 population time series. This procedure resulted in an Rmax </p><p> estimate of 1.34 which was used for both R. diaza and L. timidus. </p><p>Dispersal</p><p>The rate of dispersal between patches of suitable habitat during each time step was </p><p>185modelled with an exponential function, P = exp(Db), where D is the distance between </p><p> patch centroids and b is a constant. When D exceeds a specified maximum distance </p><p>(Dmax; set arbitrarily at a high value of 20 km), P is set to zero. The best estimate of b </p><p> was set at 2 based on observed mean and maximum dispersal distances (Hewson & Hinge</p><p>1990; Dahl & Willebrand 2005) and a sensitivity analysis examined alternative values of </p><p>1901 (less dispersal) and 3 (greater).</p><p>Stochasticity</p><p>Demographic stochasticity was implemented by sampling the number of survivors from </p><p> binomial distributions, and the number of young produced from a Poisson distribution </p><p>(Akçakaya & Root 2005). Environmental stochasticity was sampled from lognormal </p><p>195distributions. Environmental variability was correlated between populations depending on 10ESM_Changes in range limits under climate change Page 10 of 12</p><p> their spatial separation. Pairwise correlations were calculated using an exponential </p><p> function, P = exp(Db), where D is the distance between centroids of habitat patches and b</p><p> is a constant. This function was parameterised b=300, which was based on correlation-</p><p> distance relationship in annual mean temperature variation among 20 weather stations in </p><p>200the UK. Annual environmental variability for vital rates was estimated based on </p><p> variability of population sizes from time series data. ESM_Changes in range limits under climate change Page 11 of 12</p><p>Romerolagus diazi patches</p><p>Historic</p><p>201</p><p>0</p><p>205</p><p>0</p><p>208</p><p>0 ESM_Changes in range limits under climate change Page 12 of 12</p><p>205Fig. S2. Volcano rabbit habitat changes, showing size and location of habitat patches </p><p> predicted by climatic and land-use variables, with climate changing in time according to </p><p> the A2 SRES scenario. For years 2010 through 2080, the colour of grid cells indicates </p><p> habitat suitability (brighter colour more suitable), and the white outlines delineate patches</p><p> or discrete populations of the metapopulation. From 2010 to 2050, two smaller </p><p>210(northern) patches disappear, one large (western) patch splits into three smaller patches, </p><p> and two medium-sized (eastern) patches become slightly smaller. From 2050 to 2080, </p><p> two of the three western patches disappear, and the two eastern patches become </p><p> substantially smaller.</p><p>215References</p>
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