Abstract
Estimating the relative importance of habitat loss and fragmentation is necessary to estimate the potential benefits of specific management actions and to ensure that limited conservation resources are used efficiently. However, estimating relative effects is complicated because the two processes are highly correlated. Previous studies have used a wide variety of statistical methods to separate their effects and we speculated that the published results may have been influenced by the methods used. We used simulations to determine whether, under identical conditions, the following 7 methods generate different estimates of relative importance for realistically correlated landscape predictors: residual regression, model or variable selection, averaged coefficients from all supported models, summed Akaike weights, classical variance partitioning, hierarchical variance partitioning, and a multiple regression model with no adjustments for collinearity. We found that different methods generated different rankings of the predictors and that some metrics were strongly biased. Residual regression and variance partitioning were highly biased by correlations among predictors and the bias depended on the direction of a predictor’s effect (positive vs. negative). Our results suggest that many efforts to deal with the correlation between amount and fragmentation may have done more harm than good. If confounding effects are controlled and adequate thought is given to the ecological mechanisms behind modeled predictors, then standardized partial regression coefficients are unbiased estimates of the relative importance of amount and fragmentation, even when predictors are highly correlated.
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References
Barbaro L, Rossi JP, Vetillard F, Nezan J, Jactel H (2007) The spatial distribution of birds and carabid beetles in pine plantation forests: the role of landscape composition and structure. J Biogeogr 34:652–664
Bartuszevige AM, Gorchov DL, Raab L (2006) The relative importance of landscape and community features in the invasion of an exotic shrub in a fragmented landscape. Ecography 29:213–222
Belisle M, Desrochers A, Fortin MJ (2001) Influence of forest cover on the movements of forest birds: a homing experiment. Ecology 82:1893–1904
Betts MG, Forbes GJ, Diamond AW, Taylor PD (2006) Independent effects of fragmentation on songbirds in a forest mosaic. Ecol Appl 16:1076–1089
Burnham KP, Anderson (2002) Model selection and multi-model inference: a practical information–theoretic approach, 2nd edn. Springer, New York
Chevan A, Sutherland M (1991) Hierarchical partitioning. Am Stat 45:90–96
Clark AE, Troskie CG (2006) Regression and ICOMP—a simulation study. Commun Stat Simul Comput 35(3):591–603
Cohen J, Cohen P (1983) Applied multiple regression/correlation analysis for the behavioral sciences. Lawrence Erlbaum, Mahwah
Cooper CB, Walters JR (2002) Independent effects of woodland loss and fragmentation on Brown Treecreeper distribution. Biol Conserv 105:1–10
Cushmam SA, McGarrigal K (2003) Landscape-level patterns of avian diversity in the Oregon Coast Range. Ecol Monogr 73:259–281
Debuse VJ, King J, House APN (2007) Effect of fragmentation, habitat loss and within-patch habitat characteristics on ant assemblages in semi-arid woodlands of eastern Australia. Landscape Ecol 22:731–745
Dodd NL, Schweinsburg RE, Boe S (2006) Landscape-scale forest habitat relationships to tassel-eared squirrel populations: Implications for ponderosa pine forest restoration. Restor Ecol 4:537–547
Donnelly R, Marzluff JM (2006) Relative importance of habitat quantity, structure, and spatial pattern to birds in urbanizing environments. Urban Ecosyst 9:99–117
Drolet B, Desrochers A, Fortin MJ (1999) Effects of landscape structure on nesting songbird distribution in a harvested boreal forest. Condor 101:699–704
Ewers RM, Didham RK (2006) Confounding factors in the detection of species responses to habitat fragmentation. Biol Rev 81:117–142
Ewers RM, Didham RK (2007) Habitat fragmentation: panchreston or paradigm? Trends Ecol Evol 22:511
Fahrig L (1998) When does fragmentation of breeding habitat affect population survival? Ecol Modell 105:273–292
Fahrig L (2003) Effects of habitat fragmentation on biodiversity. Annu Rev Ecol Evol Syst 34:487–515
Flather CH, Bevers M (2002) Patchy reaction-diffusion and population abundance: the relative importance of habitat amount and arrangement. Am Nat 159:40–56
Fletcher RJ, Koford RR (2002) Habitat and landscape associations of breeding birds in native and restored grasslands. J Wildl Manage 66:1011–1022
Freckleton RP (2002) On the misuse of residuals in ecology: regression of residuals vs. multiple regression. J Anim Ecol 71:542–545
Gelling M, Macdonald DW, Mathews F (2007) Are hedgerows the route to increased farmland small mammal density? Use of hedgerows in British pastoral habitats. Landscape Ecol 22:1019–1032
Graham MH (2003) Confronting multicollinearity in ecological multiple regression. Ecology 84:2809–2815
Gromping U (2007) Estimators of relative importance in linear regression based on variance decomposition. Am Stat 61:139–147
Hamer TL, Flather CH, Noon BR (2006) Factors associated with grassland bird species richness: the relative roles of grassland area, landscape structure, and prey. Landscape Ecol 21:569–583
Hendrickx F, Maelfait JP, van Wingerden W, Schweiger O, Speelmans M, Aviron S, Augenstein I, Billeter R, Bailey D, Bukacek R, Burel F, Dieko¨tter T, Dirksen J, Herzog F, Liira J, Roubalova M, Vandomme V, Bugter R (2007) How landscape structure, land-use intensity and habitat diversity affect components of total arthropod diversity in agricultural landscapes. J Appl Ecol 44:340–351
Hovel KA, Lipcius RN (2001) Effects of seagrass habitat fragmentation on juvenile blue crab survival and abundance. J Exp Mar Biol Ecol 271:75–98
Koper N, Schmiegelow FKA, Merrill EH (2007) Residuals cannot distinguish between ecological effects of habitat amount and fragmentation: implications for the debate. Landscape Ecol 22:811–820
Kruskal W (1987) Relative importance by averaging over orderings. Am Stat 41:6–10
Kruskal W, Majors R (1989) Concepts of relative importance in recent scientific literature. Am Stat 43:2–6
Langlois JP, Fahrig L, Merriam G, Artsob H (2001) Landscape structure influences continental distribution of hantavirus in deer mice. Landscape Ecol 16:255–266
Legendre P, Legendre L (1998) Numerical ecology. Elsevier, Amsterdam
Lindenmayer DB, Fischer J (2007) Tackling the habitat fragmentation Panchreston. Trends Ecol Evol 22:127–132
MacNally R (2000) Regression and model-building in conservation biology, biogeography and ecology: the distinction between–and reconciliation of–‘predictive’ and ‘explanatory’ models. Biodivers Conserv 9:655–671
Magness DR, Wilkins RN, Hejl SJ (2006) Quantitative relationships among golden-cheeked warbler occurrence and landscape size, composition, and structure. Wildl Soc Bull 34:473–479
McAlpine CA, Bowen ME, Callaghan JG, Lunney D, Rhodes JR, Mitchell DL, Pullar DV, Poszingham HP (2006a) Testing alternative models for the conservation of koalas in fragmented rural-urban landscapes. Austral Ecol 31:529–544
McAlpine CA, Rhodes JR, Callaghan JG, Bowen ME, Lunney D, Mitchell DL, Pullar DV, Poszingham HP (2006b) The importance of forest area and configuration relative to local habitat factors for conserving forest mammals: a case study of koalas in Queensland, Australia. Biol Conserv 132:153–165
McGarigal K, Cushman SA (2002) Comparative evaluation of experimental approaches to the study of habitat fragmentation effects. Ecol Appl 12:335–345
McGarigal K, McComb WC (1995) Relationships between landscape structure and breeding birds in the Oregon coast range. Ecol Monogr 65:235–260
Neter J, Wasserman W, Kutner MH (1990) Applied linear statistical models, 3rd edn. Irwin, Chicago
Olson GS, Glenn EM, Anthony RG, Forsman ED, Reid JA, Loschl PJ, Ripple WJ (2004) Modeling demographic performance of northern spotted owls relative to forest habitat in Oregon. J Wildl Manage 68:1039–1053
OMNR (1998) Ontario land cover database. Ontario Ministry of Natural Resources, Peterborough Ontario
Petraitis PS, Dunham AE, Niewiarowski PH (1996) Inferring multiple causality: the limitations of path analysis. Funct Ecol 10:421–431
Radford JQ, Bennett AF (2004) Thresholds in landscape parameters: occurrence of the white-browed treecreeper Climacteris affinis in Victoria, Australia. Biol Conserv 117:375–391
Radford JQ, Bennett AF (2007) The relative importance of landscape properties for woodland birds in agricultural environments. J Appl Ecol 44:737–747
Renfrew RB, Ribic CA (2008) Multi-scale models of grassland passerine abundance in a fragmented system in Wisconsin. Landscape Ecol 23:181–193
Reunanen P, Nikula A, Monkkonen M, Hurme E, Nivala V (2002) Predicting occupancy for the Siberian flying squirrel in old-growth forest patches. Ecol Appl 12:1188–1198
Rompre G, Robinson WD, Desrochers A, Angehr G (2007) Environmental correlates of avian diversity in lowland Panama rain forests. J Biogeogr 34:802–815
Rosenberg KV, Lowe JD, Dhondt AA (1999) Effects of forest fragmentation on breeding tanagers: a continental perspective. Conserv Biol 13:568–583
Stephens SE, Rotella JJ, Lindberg MS, Taper ML, Ringelman JK (2005) Duck nest survival in-the Missouri Coteau of North Dakota: landscape effects at multiple spatial scales. Ecol Appl 15:2137–2149
Sutherland WJ, Pullin AS, Dolman PM, Knight TM (2004) The need for evidence-based conservation. Trends Ecol Evol 19:305–308
Taki H, Kevan PG, Ascher JS (2007) Landscape effects of forest loss in a pollination system. Landscape Ecol 22:1575–1587
Trzcinski MK, Fahrig L, Merriam G (1999) Independent effects of forest cover and fragmentation on the distribution of forest breeding birds. Ecol Appl 9:586–593
Turner MG (2005) Landscape ecology: what is the state of the science? Ann Rev Ecol Evol Syst 36:319–344
Villard MA, Trzcinski MK, Merriam G (1999) Fragmentation effects on forest birds: relative influence of woodland cover and configuration on landscape occupancy. Conserv Biol 13:774–783
Westphal MI, Field SA, Tyre AJ, Paton D, Possingham HP (2003) Effects of landscape pattern on bird species distribution in the Mt. Lofty Ranges, South Australia. Landscape Ecol 18:413–426
Wood PB, Bosworth SB, Dettmers R (2006) Cerulean warbler abundance and occurrence relative to large-scale edge and habitat characteristics. Condor 108:154–165
Yates MD, Muzika RM (2006) Effect of forest structure and fragmentation on site occupancy of bat species in Missouri ozark forests. J Wildl Manage 70:1238–1248
Acknowledgments
This work was funded by Natural Sciences and Engineering Research Council grants to A.C. Smith, and L. Fahrig. Members of the Friday Discussion group in the Geomatics and Landscape Ecology Research Lab provided helpful feedback on the analysis. D. Currie and two anonymous reviewers provided insightful comments on earlier drafts.
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Smith, A.C., Koper, N., Francis, C.M. et al. Confronting collinearity: comparing methods for disentangling the effects of habitat loss and fragmentation. Landscape Ecol 24, 1271–1285 (2009). https://doi.org/10.1007/s10980-009-9383-3
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DOI: https://doi.org/10.1007/s10980-009-9383-3