Abiotic factors & productivity as predictors of land cover Michael P. Simanonok INTRODUCTION Large scale spatial studies, at their core investigate the causes of landscape configuration and with land cover being the main variable to consider in regard to conservation, understanding the predictors driving landscape configuration and land cover are vital for conservation and the maintenance of biodiversity (Weins 2002). Furthermore, the inherent natural properties and production of terrestrial systems can drive the degree of human land use (Zhao et al. 2011). Protected areas, such as national parks, can be particularly vulnerable due to a marked increase in land use around the parks as well as an increase in the intensity of that use (Hansen et al. 2011). Thus, an increased understanding of what predictors determine land cover can help to identify those areas most highly at risk, as well as provide basal ecological knowledge of the determinants of natural land covers. Landscape scale spatial analysis of gross primary productivity (GPP) and net primary productivity (NPP) has become incredibly available and relatively simple to process, which allows us to ask ecologically important questions regarding relationships between ecological systems and their use (Running et al. 2004, Zhao et al. 2005). NPP and GPP are both inherently tied to precipitation, temperature, as well as elevation (Zhang et al. 2009), thus these become variables of interest. Similar work has focused primarily on land use/cover change (Parmenter et al. 2003) ¸ and has not explicitly examined if these variables can be used to directly predict land cover without considering previous land use. The question addressed herein, is what are the biotic 1 (NPP & GPP) and abiotic (precipitation, temperature, elevation) factors important in determining land use and land cover, and what is the relative importance of each? METHODS Study Area The area of analysis is the federally protected national parks within the state of Montana (Fig. 1); primarily composed of Glacier National Park, Yellowstone National Park, and Bighorn Canyon National Recreation Area (Table 1). Data for national parks were obtained from the Montana Natural Resource Information System. The rationale for the selection of protected areas is that they ideally have seen relatively little change in land use over the last ~50 years (the relevant range of our datasets) and therefore should provide a good measurement tool for how these biotic and abiotic factors determine land cover. Data These analyses require data on precipitation, temperature, elevation, NPP, GPP, and land cover. Precipitation, temperature, and elevation data were all obtained from Worldclim.org, and full methods regarding the generation of these data are described by Hijmans et al. (2005). All three of these layers are at ~1km2 resolution (30 arc second grids), and the precipitation and temperature data covered a period from 1950-2000. Monthly temperature averages from that time period for federally protected areas in Montana were averaged together to create a single mean temperature, and this was also done for the precipitation data. Estimates of NPP and GPP were derived from Moderate Resolution Imaging Spectroradiometer (MODIS) imagery (Oak Ridge National Laboratory Distributed Active Archive Center), also at 1km2 resolution. These data are generated by two NASA satellites (Terra & Aqua) that provide spectral images of the entire surface of the Earth every 1-2 days. 2 Algorithms are then run on the imaging data to provide the estimated values for NPP and GPP (Running et al. 2004, Zhao et al. 2005). Land cover data were obtained from the Multi-Resolution Land Characteristics Consortium. The National Land Cover Dataset (Fry et al. 2006) has the entire conterminous United States classified into 16 different land cover classes (Fig. 2 & Table 2) at 30m2 resolution. The data are generated from the Landsat Enhanced Thematic Mapper+ from satellite data obtained in 2006, and full methods are described by Fry et al. (2006). Using ArcMap (ESRI 2011), these data were scaled up to 1km2 resolution, so as to match the other datasets, using a resampling tool based on the composition of the nearest neighbor cells. This upscaling results in a loss of information, however given the nature of the research question as well as the fact that all other layers are at a higher resolution this resampling was found to be unavoidable. Data Manipulation The above layers (temperature, precipitation, elevation, NPP, GPP, & land cover) were imported into ArcMap (ESRI 2011) and clipped for the boundaries of the federally protected areas of Montana. The raster data were then converted into a single shapefile of point data, using the Extract Multi Values tool. After points of “no data” were removed, this resulted in 6,924 individual spatial points for the national parks, with every point including a measurement for elevation, temperature, precipitation, GPP, & NPP. Analysis The attribute table of the point data containing all variables was exported and all analyses were performed using R (R Development Core Team 2008). A regression analysis was modeled for each individual variable as predictors of land cover. Note that since a high degree of colinearity exists between these variables each regression was run separately; a full model 3 utilizing all of these potential predictors at once would not be appropriate. As a supplement for the regression analyses, an Akaike information criterion (AIC) was built comparing these individual models (Akaike 1974). This was done as an attempt to help enhance the more specific questions such as the relative importance of each climatic variable, and the appropriateness of GPP vs. NPP. RESULTS Regression analyses showed strong evidence for correlation of all three climate factors (elevation, precipitation, & temperature) with land cover type (Table 3). The p values for all three variables were so infinitesimally small that ascribing relative importance from this analysis would not be appropriate; the strong degree of colinearity between these variables is likely responsible for the similar results. Higher elevations tended to be associated with barren land and shrub/scrub land covers, with lower elevations being primarily associated with evergreen forests (Fig. 3). For precipitation, areas of high precipitation appeared to be associated with shrub/scrub cover as well, with forested areas showing lower precipitation (Fig. 4). Areas of higher temperature were visibly linked to evergreen and mixed deciduous forests, with those areas of colder temperature being more likely to have barren or more herbaceous cover (Fig. 5) Interestingly, neither GPP nor NPP were found to be important predictors of land cover (Table 3). Graphically, these results make sense, as areas with generally homogeneous trends for productivity show markedly heterogeneous land cover assemblages (Figs. 6 & 7). The AIC results for comparing the individual models did find differences in the predictive ability of the climate variables, and found precipitation to be the best fit, with elevation and temperature having relatively little difference between them (Table 4). For the 4 GPP/NPP comparison, GPP was a slightly better fit but not convincingly different, and it was a much poorer fit than any of the climate-related factors (Table 4). DISCUSSION What these results suggest is that while climatic factors may be determinants for productivity there is no observed association between productivity and land cover. Furthermore, it is likely that climate; precipitation in particular, can be a strong predictor for land cover. It is also possible that there are other factors not considered in this study that may well be able to more accurately predict land cover. With precipitation being the most important predictor found in this study, it is surprising that productivity metrics did not show strong patterns with land cover, as water is the most limiting factor for productivity in terrestrial systems (Running et al. 2004). As this study investigated land cover for federally protected areas, it is necessary to be aware of the implications for management. Given the ability to identify land cover based on spatial analysis of already existing data, it is feasible that habitat types of high concern could be easily identified and areas for future conservation be easily selected without significant time spent in the field. Similar work has been performed, such as predicting changes in land use intensity through already existing land cover models (Lambin et al. 2000). Another important consideration is the global necessity for understanding these processes in light of anthropogenic disturbance such as climate change. A major goal of data collection programs such as MODIS is for the purpose of learning about the biological processes of productivity and how climate is related; specifically investigating how ecosystems are driven and changed by humans, their innate biotic processes, and climatic determinates (Janetos & Justice 2000). Being able to predict how climate and production determine land cover provides valuable 5 utility in making specific predictions from broad climatic models, such as the Intergovernmental Panel on Climate Change’s report (IPCC 2007). Other recent broad scale spatial studies have investigated the relationship between productivity and diversity (Huston & Wolverton 2009, Belote et al. 2011) and it would be interesting to see if considering productivity and diversity of plant communities in tandem would provide accurate prediction of land cover. It has been shown that highly productive areas generally have highly diverse biotic communities (Huston & Wolverton 2009), and it could be expected that patch richness would show a similar response. The next step in addressing these questions would be to apply a similar conceptual methodology over a larger spatial area. This project, in particular, was limited in the scale of area studied due to time and computational constraints, and it would be of interest to see the results for larger areas. It is possible that at broader spatial scales, or for other systems, these patterns and relationships could differ. REFERENCES Akaike, H. 1974. A new look at the statistical model identification. IEEE Transactions on Automatic Control 19(6): 716–723. Belote, RT, Prisley, S, Jones, RH, Fitzpatrick, M, de Beurs, K. 2011. Forest productivity and tree diversity relationships depend on ecological context within mid-Atlantic and Appalachian forests (USA). Forest Ecology and Management 261: 1315-1324. ESRI 2011. ArcGIS Desktop: Release 10. Redlands, CA: Environmental Systems Research Institute. Fry, J, Xian, G, Jin, S, Dewitz, J, Homer, C, Yang, L, Barnes, C, Herold, N, and Wickham, J. 2011. Completion of the 2006 National Land Cover Database for the Conterminous United States, PE&RS 77(9): 858-864. Janetos, AC, & Justice, CO. 2000. Land cover and global productivity: a measurement strategy for the NASA programme. International Journal of Remote Sensing 21(6-7): 1491-1512. 6 Hansen, AJ, Davis, CR, Piekielek, N, Gross, J, Theobald, DM, Goetz, S, Melton, F, DeFries, R. 2011. Delineating the ecosystems containing protected areas for monitoring and management. BioScience 61: 363-373. Hijmans, RJ, Cameron, SE, Parra, JL, Jones, PG, Jarvis, A. 2005. Very high resolution interpolated climate surfaces for global land areas. International Journal of Climatology 25: 1965-1978. Huston, MA, Wolverton, S. 2009. The global distribution of net primary production: resolving the paradox. Ecological Monographs 79(3): 343-377. IPCC, 2007. Climate Change 2007: The Physical Science Basis. Contribution of Working Group I to the Fourth Assessment Report of the Intergovernmental Panel on Climate Change [Solomon, S, Qin, D, Manning, M, Chen, Z, Marquis, M, Averyt, KB, Tignor, M, & Miller, HL (eds.)]. Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA. Lambin, EF, Rounsevell, MDA, Geist, HJ. 2000. Are agricultural land-use models able to predict changes in land-use intensity? Agriculture, Ecosystems and Environment 82: 321-331. Oak Ridge National Laboratory Distributed Active Archive Center (ORNL DAAC). 2011. MODIS subsetted land products, Collection 5. Available on-line [http://daac.ornl.gov/MODIS/modis.html] from ORNL DAAC, Oak Ridge, Tennessee, U.S.A. Parmenter, AW, Hansen, A, Kennedy, RE, Cohen, W, Langner, U, Lawrence, R, Maxwell, B, Gallant, A, Aspinall, R. 2003. Land use and land cover change in the greater Yellowstone ecosystem: 1975-1995. Ecological Applications 13(3): 687-703. R Development Core Team (2008). R: A language and environment for statistical computing. R Foundation for Statistical Computing,Vienna, Austria. Running, SW, Nemani, RR, Heinsch, FA, Zhao, MS, Reeves, M, Hashimoto, H. 2004. A continuous satellite-derived measure of global terrestrial primary production. BioScience 54(6): 547-560. Wiens, JA. 2002. Central concepts and issues of landscape ecology. In: Applying Landscape Ecology in Biological Conservation Ed. K. Gutzwiller. Zhang, Y, Xu, M, Chen, H, Adams, J. 2009. Global pattern of NPP to GPP ratio derived from MODIS data: effects of ecosystem type, geographical location and climate. Global Ecology and Biogeography 18: 280-290. Zhao, MS, Heinsch, FA, Nemani, RR, Running, SW. 2005. Improvements of the MODIS terrestrial gross and net primary production global data set. Remote Sensing of Environment 95(2): 164-176. 7 Zhao, MS, Running, S, Helnsch, FA, Nemanl, R. 2011. MODIS-Derived Terrestrial Primary Production. In: B. Ramachandran et al. (eds.), Land Remote Sensing and Global Environmental Change, Remote Sensing and Digital Image Processing 11: 635-660. APPENDIX Tables Table 1: Federally protected areas within the state of Montana that were used for this analysis AREA 788194.5313 162275.3927 6449239.778 2715201.713 2403445.497 667420.3536 7749125.513 4075969323 108489779.8 640134515.2 PERIMETER 3958.46712 1806.804131 14127.06924 7287.912661 8940.00619 4083.871139 18125.18955 358956.2274 125496.0042 342100.3342 NAME Bear Paw Battlefield Fort Union Trading Post National Historic Site Grant-Kohrs Ranch National Historic Site Big Hole National Battlefield Little Bighorn Battlefield National Monument Little Bighorn Battlefield National Monument Bighorn Canyon National Recreation Area Glacier National Park Bighorn Canyon National Recreation Area Yellowstone National Park Table 2: Land cover classes as defined by the Multi-Resolution Land Characteristics Consortium. Available at www.mrlc.gov 8 ACRES 194 40 1593 670 593 164 1914 1007158 26807 158175 Table 3: Results of regression analysis; see Appendix: R Code for further information. Predictor F stat Elevation 21.99 Temperature 70.05 Precipitation 336 GPP 2.6 NPP 2.34 df 1,6922 1,6922 1,6922 1,6922 1,6922 p 2.79E-06 <2.2e-16 <2.2e-16 0.11 0.13 Table 4: Results for AIC analysis; see Appendix: R Code for further information. Predictor Elevation Temperature Precipitation GPP NPP df 3 3 3 3 3 AIC 50870.39 50822.63 50564.20 50889.76 50890.02 Delta 306.1882 258.4293 0 325.5538 325.8117 Figures Figure 1: Study area; federally protected areas within Montana. Figure 2: Land cover of study area. See Table 2 for legend. 9 Figure 3: Elevation for the study area; white areas indicate higher elevation. Range = 913 to 3167 meters. Figure 4: Mean annual precipitation from 1950-2000 for the study area; white areas indicate greater precipitation. Range = 22.6 to 68.9 millimeters. Figure 5: Mean annual temperature from 1950-2000 for the study area; white areas indicate warmer temperatures. Range = -4.4 to 9.6 °C. Figure 6: Gross Primary Productivity for the study area; white areas indicate higher productivity. Range = 52 to 1087 kg C / m2 10 Figure 7: Net Primary Productivity for the study area; white areas indicate higher productivity. Range = 42 to 740 kg C / m2 R Code > summary(alt) Call: lm(formula = lcnpsre ~ altnps, data = finaltable) Residuals: Min 1Q -43.929 -2.610 Median -1.999 3Q 7.442 Max 51.698 Coefficients: Estimate Std. Error t value Pr(>|t|) (Intercept) 46.310244 0.499020 92.802 < 2e-16 altnps -0.001285 0.000274 -4.689 2.79e-06 Residual standard error: 9.528 on 6922 degrees of freedom Multiple R-squared: 0.003167, Adjusted R-squared: 0.003023 F-statistic: 21.99 on 1 and 6922 DF, p-value: 2.794e-06 > summary(temp) Call: lm(formula = lcnpsre ~ GRID_CODE, data = finaltable) Residuals: Min 1Q -43.642 -2.924 Median -2.029 3Q 7.148 Max 51.916 Coefficients: Estimate Std. Error t value Pr(>|t|) (Intercept) 43.04197 0.16441 261.79 <2e-16 GRID_CODE 0.05038 0.00602 8.37 <2e-16 Residual standard error: 9.495 on 6922 degrees of freedom Multiple R-squared: 0.01002, Adjusted R-squared: 0.009876 F-statistic: 70.05 on 1 and 6922 DF, p-value: < 2.2e-16 > summary(prec) Call: lm(formula = lcnpsre ~ precnps, data = finaltable) Residuals: Min 1Q -44.633 -2.915 Median -1.284 3Q 6.708 Max 51.716 11 Coefficients: Estimate Std. Error t value Pr(>|t|) (Intercept) 65.05687 1.15250 56.45 <2e-16 precnps -0.37648 0.02054 -18.33 <2e-16 Residual standard error: 9.32 on 6922 degrees of freedom Multiple R-squared: 0.04629, Adjusted R-squared: 0.04615 F-statistic: 336 on 1 and 6922 DF, p-value: < 2.2e-16 > summary(gpp) Call: lm(formula = lcnpsre ~ gppre, data = finaltable) Residuals: Min 1Q -44.239 -2.236 Median -2.055 3Q 7.807 Max 51.035 Coefficients: Estimate Std. Error t value Pr(>|t|) (Intercept) 4.354e+01 3.270e-01 133.136 <2e-16 gppre 8.733e-04 5.422e-04 1.611 0.107 Residual standard error: 9.542 on 6922 degrees of freedom Multiple R-squared: 0.0003747, Adjusted R-squared: 0.0002303 F-statistic: 2.595 on 1 and 6922 DF, p-value: 0.1072 > summary(npp) Call: lm(formula = lcnpsre ~ nppre, data = finaltable) Residuals: Min 1Q -43.982 -2.227 Median -2.048 3Q 7.832 Max 51.028 Coefficients: Estimate Std. Error t value Pr(>|t|) (Intercept) 4.351e+01 3.594e-01 121.072 <2e-16 nppre 1.346e-03 8.806e-04 1.529 0.126 Residual standard error: 9.542 on 6922 degrees of freedom Multiple R-squared: 0.0003375, Adjusted R-squared: 0.0001931 F-statistic: 2.337 on 1 and 6922 DF, p-value: 0.1264 > AICc(alt,temp,prec,gpp,npp) df AIC AICc Delta weight ER Model 1 3 50870.39 50870.39 306.1882 3.251415e-67 3.075584e+66 Model 2 3 50822.63 50822.63 258.4293 7.634689e-57 1.309811e+56 Model 3 3 50564.20 50564.20 0.0000 1.000000e+00 1.000000e+00 Model 4 3 50889.75 50889.76 325.5538 2.027172e-71 4.932981e+70 Model 5 3 50890.01 50890.02 325.8117 1.781894e-71 5.612007e+70 > AICc(gpp,npp) df AIC AICc Delta weight ER Model 1 3 50889.75 50889.76 0.0000000 0.5321966 1.00000 Model 2 3 50890.01 50890.02 0.2579299 0.4678034 1.13765 > AICc(alt,temp,prec) df AIC AICc Delta weight ER Model 1 3 50870.39 50870.39 306.1882 3.251415e-67 3.075584e+66 Model 2 3 50822.63 50822.63 258.4293 7.634689e-57 1.309811e+56 Model 3 3 50564.20 50564.20 0.0000 1.000000e+00 1.000000e+00 12