Abstract
Motivated by the need to solve ecological problems (climate change, habitat fragmentation and biological invasions), there has been increasing interest in species distribution models (SDMs). Predictions from these models inform conservation policy, invasive species management and disease-control measures. However, predictions are subject to uncertainty, the degree and source of which is often unrecognized. Here, we review the SDM literature in the context of uncertainty, focusing on three main classes of SDM: niche-based models, demographic models and process-based models. We identify sources of uncertainty for each class and discuss how uncertainty can be minimized or included in the modelling process to give realistic measures of confidence around predictions. Because this has typically not been performed, we conclude that uncertainty in SDMs has often been underestimated and a false precision assigned to predictions of geographical distribution. We identify areas where development of new statistical tools will improve predictions from distribution models, notably the development of hierarchical models that link different types of distribution model and their attendant uncertainties across spatial scales. Finally, we discuss the need to develop more defensible methods for assessing predictive performance, quantifying model goodness-of-fit and for assessing the significance of model covariates.
Footnotes
References
- 1
Gaston K. J.& Blackburn T. M. . 1999A critique for macroecology. Oikos 84, 353–368.doi:10.2307/3546417 (doi:10.2307/3546417). Crossref, ISI, Google Scholar - 2
Kearney M., Porter W. P., Williams C., Ritchie S.& Hoffmann A. A. . 2009Integrating biophysical models and evolutionary theory to predict climatic impacts on species' ranges: the dengue mosquito Aedes aegypti in Australia. Funct. Ecol. 23, 528–538.doi:10.1111/j.1365-2435.2008.01538.x (doi:10.1111/j.1365-2435.2008.01538.x). Crossref, ISI, Google Scholar - 3
Rouget M., Richardson D. M., Milton S. J.& Polakow D. . 2001Predicting invasion dynamics of four alien Pinus species in a highly fragmented semi-arid shrubland in South Africa. Plant Ecol. 152, 79–92.doi:10.1023/A:1011412427075 (doi:10.1023/A:1011412427075). Crossref, ISI, Google Scholar - 4
Loiselle B. A., Howell C. A., Graham C. H., Goerck J. M., Brooks T., Smith K. G.& Williams P. H. . 2003Avoiding pitfalls of using species distribution models in conservation planning. Conserv. Biol. 17, 1591–1600.doi:10.1111/j.1523-1739.2003.00233.x (doi:10.1111/j.1523-1739.2003.00233.x). Crossref, ISI, Google Scholar - 5
Rodríguez J. P., Brotons L., Bustamante J.& Seoane J. . 2007The application of predictive modelling of species distribution to biodiversity conservation. Div. Distrib. 13, 243–251.doi:10.1111/j.1472-4642.2007.00356.x (doi:10.1111/j.1472-4642.2007.00356.x). Crossref, ISI, Google Scholar - 6
Anderson B. J., 2009Using distribution models to test alternative hypotheses about a species' environmental limits and recovery prospects. Biol. Conserv. 142, 488–499.doi:10.1016/j.biocon.2008.10.036 (doi:10.1016/j.biocon.2008.10.036). Crossref, ISI, Google Scholar - 7
Parmesan C.& Yohe G. . 2003A globally coherent fingerprint of climate change impacts across natural systems. Nature 421, 37–42.doi:10.1038/nature01286 (doi:10.1038/nature01286). Crossref, PubMed, ISI, Google Scholar - 8
Kremen C., 2008Aligning conservation priorities across taxa in Madagascar with high-resolution planning tools. Science 320, 222.doi:10.1126/science.1155193 (doi:10.1126/science.1155193). Crossref, PubMed, ISI, Google Scholar - 9
Vaughan I. P.& Ormerod S. J. . 2003Improving the quality of distribution models for conservation by addressing shortcomings in the field collection of training data. Conserv. Biol. 17, 1601–1611.doi:10.1111/j.1523-1739.2003.00359.x (doi:10.1111/j.1523-1739.2003.00359.x). Crossref, ISI, Google Scholar - 10
Huntley B., Collingham Y. C., Willis S. G.& Green R. E. . 2008Potential impacts of climatic change on European breeding birds. PLoS ONE 3, e1439.doi:10.1371/journal.pone.0001439 (doi:10.1371/journal.pone.0001439). Crossref, PubMed, ISI, Google Scholar - 11
Julliard R., Jiguet F.& Couvet D. . 2004Common birds facing global changes: what makes a species at risk?Glob. Change Biol. 10, 148–154.doi:10.1111/j.1365-2486.2003.00723.x (doi:10.1111/j.1365-2486.2003.00723.x). Crossref, ISI, Google Scholar - 12
Araújo M. B.& Guisan A. . 2006Five (or so) challenges for species distribution modelling. J. Biogeogr. 33, 1677–1688.doi:10.1111/j.1365-2699.2006.01584.x (doi:10.1111/j.1365-2699.2006.01584.x). Crossref, ISI, Google Scholar - 13
Heikkinen R. K., Luoto M., Araujo M. B., Virkkala R., Thuiller W.& Sykes M. T. . 2006Methods and uncertainties in bioclimatic envelope modelling under climate change. Prog. Phys. Geogr. 30, 751–777.doi:10.1177/0309133306071957 (doi:10.1177/0309133306071957). Crossref, ISI, Google Scholar - 14
Thuiller W. . 2004Patterns and uncertainties of species' range shifts under climate change. Glob. Change Biol. 10, 2020–2027.doi:10.1111/j.1365-2486.2004.00859.x (doi:10.1111/j.1365-2486.2004.00859.x). Crossref, ISI, Google Scholar - 15
Buisson L., Thuiller W., Casajus N., Lek S.& Grenouillet G. . 2010Uncertainty in ensemble forecasting of species distribution. Glob. Change Biol. 16, 1145–1157.doi:10.1111/j.1365-2486.2009.02000.x (doi:10.1111/j.1365-2486.2009.02000.x). Crossref, ISI, Google Scholar - 16
Thomas C. D., 2004Extinction risk from climate change. Nature 427, 145–148.doi:10.1038/nature02121 (doi:10.1038/nature02121). Crossref, PubMed, ISI, Google Scholar - 17
Berkhout F. . 2010Reconstructing boundaries and reason in the climate debate. Glob. Environ. Change 20, 565–569.doi:10.1016/j.gloenvcha.2010.07.006 (doi:10.1016/j.gloenvcha.2010.07.006). Crossref, ISI, Google Scholar - 18
Raiffa H.& Schlaifer R. . 1968Applied statistical decision theory.Chichester, UK: Wiley. Google Scholar - 19
Austin M. P. . 2002Spatial prediction of species distribution: an interface between ecological theory and statistical modelling. Ecol. Model. 157, 101–118.doi:10.1016/S0304-3800(02)00205-3 (doi:10.1016/S0304-3800(02)00205-3). Crossref, ISI, Google Scholar - 20
Elith J., 2006Novel methods improve prediction of species' distributions from occurrence data. Ecography 29, 129–151.doi:10.1111/j.2006.0906-7590.04596.x (doi:10.1111/j.2006.0906-7590.04596.x). Crossref, ISI, Google Scholar - 21
Elith J.& Leathwick J. R. . 2009Species distribution models: ecological explanation and prediction across space and time. Annu. Rev. Ecol. Evol. Syst. 40, 677–697.doi:10.1146/annurev.ecolsys.110308.120159 (doi:10.1146/annurev.ecolsys.110308.120159). Crossref, ISI, Google Scholar - 22
Guisan A.& Thuiller W. . 2005Predicting species distribution: offering more than simple habitat models. Ecol. Lett. 8, 993–1009.doi:10.1111/j.1461-0248.2005.00792.x (doi:10.1111/j.1461-0248.2005.00792.x). Crossref, ISI, Google Scholar - 23
Hernandez P. A., Graham C. H., Master L. L.& Albert D. L. . 2006The effect of sample size and species characteristics on performance of different species distribution modeling methods. Ecography 29, 773–785.doi:10.1111/j.0906-7590.2006.04700.x (doi:10.1111/j.0906-7590.2006.04700.x). Crossref, ISI, Google Scholar - 24
Pearson R. G., Raxworthy C. J., Nakamura M.& Peterson A. T. . 2007Predicting species distributions from small numbers of occurrence records: a test case using cryptic geckos in Madagascar. J. Biogeogr. 34, 102–117.doi:10.1111/j.1365-2699.2006.01594.x (doi:10.1111/j.1365-2699.2006.01594.x). Crossref, ISI, Google Scholar - 25
Pulliam H. R. . 2000On the relationship between niche and distribution. Ecol. Lett. 3, 349–361.doi:10.1046/j.1461-0248.2000.00143.x (doi:10.1046/j.1461-0248.2000.00143.x). Crossref, ISI, Google Scholar - 26
Alstroem P. E. R., Davidson P., Duckworth J. W., Eames J. C., Le T. T., Nguyen C., Olsson U., Robson C.& Timmins R. . 2010Description of a new species of Phylloscopus warbler from Vietnam and Laos. Ibis 152, 145–168.doi:10.1111/j.1474-919X.2009.00990.x (doi:10.1111/j.1474-919X.2009.00990.x). Crossref, ISI, Google Scholar - 27
Ceballos G.& Ehrlich P. R. . 2009Discoveries of new mammal species and their implications for conservation and ecosystem services. Proc. Natl Acad. Sci. USA 106, 3841.doi:10.1073/pnas.0812419106 (doi:10.1073/pnas.0812419106). Crossref, PubMed, ISI, Google Scholar - 28
Eriksson O. . 1996Regional dynamics of plants: a review of evidence for remnant, source–sink and metapopulations. Oikos 77, 248–258.doi:10.2307/3546063 (doi:10.2307/3546063). Crossref, ISI, Google Scholar - 29
Kawecki T. J. . 1995Demography of source—sink populations and the evolution of ecological niches. Evol. Ecol. 9, 38–44.doi:10.1007/BF01237695 (doi:10.1007/BF01237695). Crossref, ISI, Google Scholar - 30
Dennis R. L., Sparks T. H.& Hardy P. B. . 1999Bias in butterfly distribution maps: the effects of sampling effort. J. Insect Conserv. 3, 33–42.doi:10.1023/A:1009678422145 (doi:10.1023/A:1009678422145). Crossref, ISI, Google Scholar - 31
Hortal J., Jiménez-Valverde A., Gómez J. F., Lobo J. M.& Baselga A. . 2008Historical bias in biodiversity inventories affects the observed environmental niche of the species. Oikos 117, 847–858.doi:10.1111/j.0030-1299.2008.16434.x (doi:10.1111/j.0030-1299.2008.16434.x). Crossref, ISI, Google Scholar - 32
Prendergast J. R., Wood S. N., Lawton J. H.& Eversham B. C. . 1993Correcting for variation in recording effort in analyses of diversity hotspots. Biodivers. Lett. 1, 39–53.doi:10.2307/2999649 (doi:10.2307/2999649). Crossref, Google Scholar - 33
Guisan A.& Zimmermann N. E. . 2000Predictive habitat distribution models in ecology. Ecol. Model. 135, 147–186.doi:10.1016/S0304-3800(00)00354-9 (doi:10.1016/S0304-3800(00)00354-9). Crossref, ISI, Google Scholar - 34
Johnson G. L., Daly C., Taylor G. H.& Hanson C. L. . 2010Spatial variability and interpolation of stochastic weather simulation model parameters. J. Appl. Meteorol. 39, 778–796.doi:10.1175/1520-0450(2000)039<0778:SVAIOS>2.0.CO;2 (doi:10.1175/1520-0450(2000)039<0778:SVAIOS>2.0.CO;2). Crossref, ISI, Google Scholar - 35
Vetaas O. R. . 2002Realized and potential climate niches: a comparison of four Rhododendron tree species. J. Biogeogr. 29, 545–554.doi:10.1046/j.1365-2699.2002.00694.x (doi:10.1046/j.1365-2699.2002.00694.x). Crossref, ISI, Google Scholar - 36
Malanson G. P., Westman W. E.& Yan Y. L. . 1992Realized versus fundamental niche functions in a model of chaparral response to climatic change. Ecol. Model. 64, 261–277.doi:10.1016/0304-3800(92)90026-B (doi:10.1016/0304-3800(92)90026-B). Crossref, ISI, Google Scholar - 37
Soberón J. . 2007Grinnellian and Eltonian niches and geographic distributions of species. Ecol. Lett. 10, 1115–1123.doi:10.1111/j.1461-0248.2007.01107.x (doi:10.1111/j.1461-0248.2007.01107.x). Crossref, PubMed, ISI, Google Scholar - 38
Case T. J.& Taper M. L. . 2000Interspecific competition, environmental gradients, gene flow, and the coevolution of species' borders. Am. Nat. 155, 583–605.doi:10.1086/303351 (doi:10.1086/303351). Crossref, PubMed, ISI, Google Scholar - 39
Secondi J., Bretagnolle V., Compagnon C.& Faivre B. . 2003Species-specific song convergence in a moving hybrid zone between two passerines. Biol. J. Linnean Soc. 80, 507–517.doi:10.1046/j.1095-8312.2003.00248.x (doi:10.1046/j.1095-8312.2003.00248.x). Crossref, ISI, Google Scholar - 40
Pearson R. G.& Dawson T. P. . 2004Bioclimate envelope models: what they detect and what they hide—response to Hampe (2004). Glob. Ecol. Biogeogr. 13, 471–473.doi:10.1111/j.1466-822X.2004.00112.x (doi:10.1111/j.1466-822X.2004.00112.x). Crossref, Google Scholar - 41
Hampe A. . 2004Bioclimate envelope models: what they detect and what they hide. Glob. Ecol. Biogeogr. 13, 469–471.doi:10.1111/j.1466-822X.2004.00090.x (doi:10.1111/j.1466-822X.2004.00090.x). Crossref, Google Scholar - 42
Peterson A. T., 2009The climate envelope may not be empty. Proc. Natl Acad. Sci. USA 106, E47.doi:10.1073/pnas.0809722106 (doi:10.1073/pnas.0809722106). Crossref, PubMed, ISI, Google Scholar - 43
Jiménez-Valverde A., Barve N., Lira-Noriega A., Maher S. P., Nakazawa Y., Papeş M., Soberón J., Sukumaran J.& Peterson A. P. . 2011Dominant climate influences on North American bird distributions. Glob. Ecol. Biogeogr. 20, 114–118.doi:10.1111/j.1466-8238.2010.00574.x (doi:10.1111/j.1466-8238.2010.00574.x). Crossref, Google Scholar - 44
Beale C. M., Lennon J. J., Yearsley J. M., Brewer M. J.& Elston D. A. . 2010Regression analysis of spatial data. Ecol. Lett. 13, 246–264.doi:10.1111/j.1461-0248.2009.01422.x (doi:10.1111/j.1461-0248.2009.01422.x). Crossref, PubMed, ISI, Google Scholar - 45
Beaumont L. J., Hughes L.& Poulsen M. . 2005Predicting species distributions: use of climatic parameters in BIOCLIM and its impact on predictions of species' current and future distributions. Ecol. Model. 186, 251–270.doi:10.1016/j.ecolmodel.2005.01.030 (doi:10.1016/j.ecolmodel.2005.01.030). Crossref, ISI, Google Scholar - 46
Carpenter G., Gillison A. N.& Winter J. . 1993DOMAIN: a flexible modelling procedure for mapping potential distributions of plants and animals. Biodivers. Conserv. 2, 667–680.doi:10.1007/BF00051966 (doi:10.1007/BF00051966). Crossref, ISI, Google Scholar - 47
McMahon S. M., 2011Improving assessment and modelling of climate change impacts on global terrestrial biodiversity. Trends Ecol. Evol. 26, 249–259.doi:10.1016/j.tree.2011.02.012 (doi:10.1016/j.tree.2011.02.012). Crossref, PubMed, ISI, Google Scholar - 48
Vaughan I. P.& Ormerod S. J. . 2005The continuing challenges of testing species distribution models. J. Appl. Ecol. 42, 720–730.doi:10.1111/j.1365-2664.2005.01052.x (doi:10.1111/j.1365-2664.2005.01052.x). Crossref, ISI, Google Scholar - 49
Guisan A., Lehmann A., Ferrier S., Austin M., Overton J., Aspinall R.& Hastie T. . 2006Making better biogeographical predictions of species' distributions. J. Appl. Ecol. 43, 386–392.doi:10.1111/j.1365-2664.2006.01164.x (doi:10.1111/j.1365-2664.2006.01164.x). Crossref, ISI, Google Scholar - 50
Lobo J. M., Jiménez-Valverde A.& Real R. . 2008AUC: a misleading measure of the performance of predictive distribution models. Glob. Ecol. Biogeogr. 17, 145–151.doi:10.1111/j.1466-8238.2007.00358.x (doi:10.1111/j.1466-8238.2007.00358.x). Crossref, Google Scholar - 51
Beale C. M., Lennon J. J.& Gimona A. . 2009European bird distributions still show few climate associations. Proc. Natl Acad. Sci. USA 106, E41–E43.doi:10.1073/pnas.0902229106 (doi:10.1073/pnas.0902229106). Crossref, ISI, Google Scholar - 52
Stockwell D. R.& Peterson A. T. . 2002Effects of sample size on accuracy of species distribution models. Ecol. Model. 148, 1–13.doi:10.1016/S0304-3800(01)00388-X (doi:10.1016/S0304-3800(01)00388-X). Crossref, ISI, Google Scholar - 53
Murphy J. M., Sexton D. M., Barnett D. N., Jones G. S., Webb M. J., Collins M.& Stainforth D. A. . 2004Quantification of modelling uncertainties in a large ensemble of climate change simulations. Nature 430, 768–772.doi:10.1038/nature02771 (doi:10.1038/nature02771). Crossref, PubMed, ISI, Google Scholar - 54
Heikkinen R. K., Luoto M., Virkkala R., Pearson R. G.& Körber J. H. . 2007Biotic interactions improve prediction of boreal bird distributions at macro-scales. Glob. Ecol. Biogeogr. 16, 754–763.doi:10.1111/j.1466-8238.2007.00345.x (doi:10.1111/j.1466-8238.2007.00345.x). Crossref, Google Scholar - 55
Williams J. W., Jackson S. T.& Kutzbach J. E. . 2007Projected distributions of novel and disappearing climates by 2100 AD. Proc. Natl Acad. Sci. USA 104, 5738.doi:10.1073/pnas.0606292104 (doi:10.1073/pnas.0606292104). Crossref, PubMed, ISI, Google Scholar - 56
Royle J. A.& Link W. A. . 2006Generalized site occupancy models allowing for false positive and false negative errors. Ecology 87, 835–841.doi:10.1890/0012-9658(2006)87[835:GSOMAF]2.0.CO;2 (doi:10.1890/0012-9658(2006)87[835:GSOMAF]2.0.CO;2). Crossref, PubMed, ISI, Google Scholar - 57
Royle J. A., Kery M., Gautier R.& Schmid H. . 2007Hierarchical spatial models of abundance and occurrence from imperfect survey data. Ecol. Monogr. 77, 465–481.doi:10.1890/06-0912.1 (doi:10.1890/06-0912.1). Crossref, ISI, Google Scholar - 58
Draper D. . 1995Assessment and propagation of model uncertainty. J. R. Stat. Soc. B 57, 45–97. Google Scholar - 59
Faisal A., Dondelinger F., Husmeier D.& Beale C. M. . 2010Inferring species interaction networks from species abundance data: a comparative evaluation of various statistical and machine learning methods. Ecol. Inform. 5, 451–464.doi:10.1016/j.ecoinf.2010.06.005 (doi:10.1016/j.ecoinf.2010.06.005). Crossref, ISI, Google Scholar - 60
Milns I., Beale C. M.& Smith V. A. . 2010Revealing ecological networks using Bayesian network inference algorithms. Ecology 91, 1892–1899.doi:10.1890/09-0731.1 (doi:10.1890/09-0731.1). Crossref, PubMed, ISI, Google Scholar - 61
- 62
Wilson R. J., Davies Z. G.& Thomas C. D. . 2010Linking habitat use to range expansion rates in fragmented landscapes: a metapopulation approach. Ecography 33, 73–82.doi:10.1111/j.1600-0587.2009.06038.x (doi:10.1111/j.1600-0587.2009.06038.x). Crossref, ISI, Google Scholar - 63
Sillett T. S., Holmes R. T.& Sherry T. W. . 2000Impacts of a global climate cycle on population dynamics of a migratory songbird. Science 288, 2040.doi:10.1126/science.288.5473.2040 (doi:10.1126/science.288.5473.2040). Crossref, PubMed, ISI, Google Scholar - 64
Pearce-Higgins J. W., Yalden D. W., Dougall T. W.& Beale C. M. . 2009Does climate change explain the decline of a trans-Saharan Afro-Palaearctic migrant?Oecologia 159, 649–659.doi:10.1007/s00442-008-1242-4 (doi:10.1007/s00442-008-1242-4). Crossref, PubMed, ISI, Google Scholar - 65
Pearce-Higgins J. W., Dennis P., Whittingham M. J.& Yalden D. W. . 2010Impacts of climate on prey abundance account for fluctuations in a population of a northern wader at the southern edge of its range. Glob. Change Biol. 16, 12–23.doi:10.1111/j.1365-2486.2009.01883.x (doi:10.1111/j.1365-2486.2009.01883.x). Crossref, ISI, Google Scholar - 66
Both C., Bouwhuis S., Lessells C. M.& Visser M. E. . 2006Climate change and population declines in a long-distance migratory bird. Nature 441, 81–83.doi:10.1038/nature04539 (doi:10.1038/nature04539). Crossref, PubMed, ISI, Google Scholar - 67
Ovaskainen O.& Hanski I. . 2004From individual behavior to metapopulation dynamics: unifying the patchy population and classic metapopulation models. Am. Nat. 164, 364–377.doi:10.1086/423151 (doi:10.1086/423151). Crossref, PubMed, ISI, Google Scholar - 68
Griffiths R. A., Sewell D.& McCrea R. S. . 2010Dynamics of a declining amphibian metapopulation: survival, dispersal and the impact of climate. Biol. Conserv. 143, 485–491.doi:10.1016/j.biocon.2009.11.017 (doi:10.1016/j.biocon.2009.11.017). Crossref, ISI, Google Scholar - 69
McLaughlin J. F., Hellmann J. J., Boggs C. L.& Ehrlich P. R. . 2002Climate change hastens population extinctions. Proc. Natl Acad. Sci. USA 99, 6070.doi:10.1073/pnas.052131199 (doi:10.1073/pnas.052131199). Crossref, PubMed, ISI, Google Scholar - 70
Berger D., Walters R.& Gotthard K. . 2008What limits insect fecundity? Body size-and temperature-dependent egg maturation and oviposition in a butterfly. Funct. Ecol. 22, 523–529.doi:10.1111/j.1365-2435.2008.01392.x (doi:10.1111/j.1365-2435.2008.01392.x). Crossref, ISI, Google Scholar - 71
Baguette M. . 2004The classical metapopulation theory and the real, natural world: a critical appraisal. Basic Appl. Ecol. 5, 213–224.doi:10.1016/j.baae.2004.03.001 (doi:10.1016/j.baae.2004.03.001). Crossref, ISI, Google Scholar - 72
Hanski I. . 2004Metapopulation theory, its use and misuse. Basic Appl. Ecol. 5, 225–229.doi:10.1016/j.baae.2004.03.002 (doi:10.1016/j.baae.2004.03.002). Crossref, ISI, Google Scholar - 73
Stillman R.& Brown A. F. . 1998Pattern in the distribution of Britain's upland breeding birds. J. Biogeogr. 25, 73–82.doi:10.1046/j.1365-2699.1998.251169.x (doi:10.1046/j.1365-2699.1998.251169.x). Crossref, ISI, Google Scholar - 74
Cattadori I. M., Haydon D. T.& Hudson P. J. . 2005Parasites and climate synchronize red grouse populations. Nature 433, 737–741.doi:10.1038/nature03276 (doi:10.1038/nature03276). Crossref, PubMed, ISI, Google Scholar - 75
Lindström Å.& Agrell J. . 1999Global change and possible effects on the migration and reproduction of Arctic-breeding waders. Ecol. Bull. 47, 145–159. Google Scholar - 76
Moss R., Oswald J.& Baines D. . 2001Climate change and breeding success: decline of the capercaillie in Scotland. J. Anim. Ecol. 70, 47–61.doi:10.1046/j.1365-2656.2001.00473.x (doi:10.1046/j.1365-2656.2001.00473.x). Crossref, ISI, Google Scholar - 77
Willis S. G., Hill J. K., Thomas C. D., Roy D. B., Fox R., Blakeley D. S.& Huntley B. . 2009Assisted colonization in a changing climate: a test-study using two UK butterflies. Conserv. Lett. 2, 46–52.doi:10.1111/j.1755-263X.2008.00043.x (doi:10.1111/j.1755-263X.2008.00043.x). Crossref, ISI, Google Scholar - 78
Morin X., Viner D.& Chuine I. . 2008Tree species range shifts at a continental scale: new predictive insights from a process-based model. J. Ecol. 96, 784–794.doi:10.1111/j.1365-2745.2008.01369.x (doi:10.1111/j.1365-2745.2008.01369.x). Crossref, ISI, Google Scholar - 79
Kearney M.& Porter W. . 2009Mechanistic niche modelling: combining physiological and spatial data to predict species' ranges. Ecol. Lett. 12, 334–350.doi:10.1111/j.1461-0248.2008.01277.x (doi:10.1111/j.1461-0248.2008.01277.x). Crossref, PubMed, ISI, Google Scholar - 80
Kearney M. R., Wintle B. A.& Porter W. P. . 2010Correlative and mechanistic models of species distribution provide congruent forecasts under climate change. Conserv. Lett. 3, 203–213.doi:10.1111/j.1755-263X.2010.00097.x (doi:10.1111/j.1755-263X.2010.00097.x). Crossref, ISI, Google Scholar - 81
Kearney M., Phillips B. L., Tracy C. R., Christian K. A., Betts G.& Porter W. P. . 2008Modelling species distributions without using species distributions: the cane toad in Australia under current and future climates. Ecography 31, 423–434.doi:10.1111/j.0906-7590.2008.05457.x (doi:10.1111/j.0906-7590.2008.05457.x). Crossref, ISI, Google Scholar - 82
Keenan T., MariaSerra J., Lloret F., Ninyerola M.& Sabate S. . 2011Predicting the future of forests in the Mediterranean under climate change, with niche-and process-based models: CO2 matters!. Glob. Change Biol. 17, 565–579.doi:10.1111/j.1365-2486.2010.02254.x (doi:10.1111/j.1365-2486.2010.02254.x). Crossref, ISI, Google Scholar - 83
McInerny G. J., Turner J. R. G., Wong H. Y., Travis J. M. J.& Benton T. G. . 2009How range shifts induced by climate change affect neutral evolution. Proc. R. Soc. B 276, 1527.doi:10.1098/rspb.2008.1567 (doi:10.1098/rspb.2008.1567). Link, ISI, Google Scholar - 84
Bonan G. B., Levis S., Sitch S., Vertenstein M.& Oleson K. W. . 2003A dynamic global vegetation model for use with climate models: concepts and description of simulated vegetation dynamics. Glob. Change Biol. 9, 1543–1566.doi:10.1046/j.1365-2486.2003.00681.x (doi:10.1046/j.1365-2486.2003.00681.x). Crossref, ISI, Google Scholar - 85
Sitch S., 2003Evaluation of ecosystem dynamics, plant geography and terrestrial carbon cycling in the LPJ dynamic global vegetation model. Glob. Change Biol. 9, 161–185.doi:10.1046/j.1365-2486.2003.00569.x (doi:10.1046/j.1365-2486.2003.00569.x). Crossref, ISI, Google Scholar - 86
Fisher R., McDowell N., Purves D., Moorcroft P., Sitch S., Cox P., Huntingford C., Meir P.& Woodward F. I. . 2010Assessing uncertainties in a second-generation dynamic vegetation model caused by ecological scale limitations. New Phytol. 187, 666–681.doi:10.1111/j.1469-8137.2010.03340.x (doi:10.1111/j.1469-8137.2010.03340.x). Crossref, PubMed, ISI, Google Scholar - 87
Belsky A. J. . 1990Tree/grass ratios in East African savannas: a comparison of existing models. J. Biogeogr. 17, 483–489.doi:10.2307/2845380 (doi:10.2307/2845380). Crossref, ISI, Google Scholar - 88
Neilson R. P., Pitelka L. F., Solomon A. M., Nathan R., Midgley G. F., Fragoso J. M., Lischke H.& Thompson K. . 2005Forecasting regional to global plant migration in response to climate change. Bioscience 55, 749–759.doi:10.1641/0006-3568(2005)055[0749:FRTGPM]2.0.CO;2 (doi:10.1641/0006-3568(2005)055[0749:FRTGPM]2.0.CO;2). Crossref, ISI, Google Scholar - 89
Rosenthal R. . 1979The file drawer problem and tolerance for null results. Psychol. Bull. 86, 638–641.doi:10.1037/0033-2909.86.3.638 (doi:10.1037/0033-2909.86.3.638). Crossref, ISI, Google Scholar - 90
Rivington M., Matthews K. B., Bellocchi G.& Buchan K. . 2006Evaluating uncertainty introduced to process-based simulation model estimates by alternative sources of meteorological data. Agric. Syst. 88, 451–471.doi:10.1016/j.agsy.2005.07.004 (doi:10.1016/j.agsy.2005.07.004). Crossref, ISI, Google Scholar - 91
Duncan R. P., Cassey P.& Blackburn T. M. . 2009Do climate envelope models transfer? A manipulative test using dung beetle introductions. Proc. R. Soc. B 276, 1449–1457.doi:10.1098/rspb.2008.1801 (doi:10.1098/rspb.2008.1801). Link, ISI, Google Scholar - 92
Randin C. F., Dirnböck T., Dullinger S., Zimmermann N. E., Zappa M.& Guisan A. . 2006Are niche-based species distribution models transferable in space?J. Biogeogr. 33, 1689–1703.doi:10.1111/j.1365-2699.2006.01466.x (doi:10.1111/j.1365-2699.2006.01466.x). Crossref, ISI, Google Scholar - 93
Tingley M. W., Monahan W. B., Beissinger S. R.& Moritz C. . 2009Birds track their Grinnellian niche through a century of climate change. Proc. Natl Acad. Sci. USA 106, 19 637–19 643.doi:10.1073/pnas.0901562106 (doi:10.1073/pnas.0901562106). Crossref, ISI, Google Scholar - 94
Araújo M. B.& New M. . 2007Ensemble forecasting of species distributions. Trends Ecol. Evol. 22, 42–47.doi:10.1016/j.tree.2006.09.010 (doi:10.1016/j.tree.2006.09.010). Crossref, PubMed, ISI, Google Scholar - 95
Phillips S. J.& Dudik M. . 2008Modeling of species distributions with Maxent: new extensions and a comprehensive evaluation. Ecography 31, 161–175.doi:10.1111/j.0906-7590.2008.5203.x (doi:10.1111/j.0906-7590.2008.5203.x). Crossref, ISI, Google Scholar - 96
Wood A. W., Leung L. R., Sridhar V.& Lettenmaier D. P. . 2004Hydrologic implications of dynamical and statistical approaches to downscaling climate model outputs. Clim. Change 62, 189–216.doi:10.1023/B:CLIM.0000013685.99609.9e (doi:10.1023/B:CLIM.0000013685.99609.9e). Crossref, ISI, Google Scholar - 97
Anderson B. J., Akcakaya H. R., Araujo M. B., Fordham D. A., Martinez-Meyer E., Thuiller W.& Brook B. W. . 2009Dynamics of range margins for metapopulations under climate change. Proc. R. Soc. B 276, 1415.doi:10.1098/rspb.2008.1681 (doi:10.1098/rspb.2008.1681). Link, ISI, Google Scholar - 98
Possingham H. P. . 1996Decision theory and biodiversity management: how to manage a metapopulation. Frontiers of population ecology (eds, Floyd R. B., Sheppard A. W.& de Barro P. J. ), pp. 391–398. Melbourne, Australia: CSIRO Publishing. Google Scholar - 99
Possingham H. . 2010The business of biodiversity: applying decision theory principles to nature conservation.Fitzroy, Australia: Australian Conservation Foundation. Google Scholar - 100
Regan H. M., Ben-Haim Y., Langford B., Wilson W. G., Lundberg P., Andelman S. J.& Burgman M. A. . 2005Robust decision-making under severe uncertainty for conservation management. Ecol. Appl. 15, 1471–1477.doi:10.1890/03-5419 (doi:10.1890/03-5419). Crossref, ISI, Google Scholar


