The challenge of statistically identifying species-resource relationships on an uncooperative landscape Or… Facts, true facts, and statistics: a lesson in numeracy Barry D. Smith & Kathy Martin Canadian Wildlife Service, Pacific Wildlife Research Centre Delta, B.C., Canada Clive Goodinson Free Agent,Vancouver, B.C., Canada Species-Habitat Associations Objective: To incorporate habitat suitability predictions into a stand-level forest ecosystem model + = Can we show statistically that the relative quantity of a resource on the landscape predicts the presence of a species such as Northern Flicker? Logistic regression model output Predicted 0 1 Predicted 0 1 0 123 16 1 9 74 Observed Logistic regression model Observed Groups and Predicted Probabilities 20 + 1 + I 1 I I 1 I F I 1 1 I R 15 + 1 1 + E I 1 1 1 1 I Q I 1 1 1 111 1 1 I U I 11 11 11 111 1 11 I E 10 + 1 11111 11 11111 11 1 + N I 1 1 10111101 11111111 1 I C I 011110011001110101111 1 1 I Y I 01110000100111000111111 1 I 5 + 00 001100000000110000001111111 11 + I 001000100000000000000001111101 1 11 I I 0 00000000000000000000000010001000110 11 I I 0 1 000000000000000000000000001000000000011011 11 1 I Predicted --------------+--------------+--------------+--------------Prob: 0 .25 .5 .75 1 Group: 000000000000000000000000000000111111111111111111111111111111 0 = Absent 1 = Present Predicted 0 Observed 1 0 1 Habitat is over 100% saturated; birds occur in areas of poor habitat. Sampling intensity is too low; birds occur within good habitat but sampling does not capture all occurrences. Habitat is not 100% saturated; there are areas of good habitat which are unoccupied. Spatial variability is too low or spatial periodicity of key habitat attributes is too high, given sampling intensity. The playback tape pulls in individuals from outside the point-count radius. So, can we expect be successful in detecting species-habitat associations when they exist? We use simulations where: we generated a landscape, then • populated that landscape with a (territorial) species, then • sampled the species and landscape repeatedly to assess our ability to detect a known association Sample Simulation > Sample Sim’on To be as realistic as possible we need to make decisions concerning… •The characteristics of the landscape (resources) •The species’ distribution on thelandscape • The sampling method • The statistical model(s) Spatial contrast is essential for, but doesn’t guarantee, success High Landscape Spatial Periodicity (SP) Medium Landscape Spatial Periodicity (SP) Low Landscape Spatial Periodicity (SP) It might help to conceptualize required resources by consolidating them into four fundamental suites: • Shelter (e.g., sleeping, breeding) • Food (self, provisioning) • Comfort (e.g. weather, temperature) • Safety (predation risk) To be as realistic as possible we had to make decisions concerning: •The characteristics of the landscape •The species’ distribution on thelandscape • The sampling method • The statistical model(s) Territory establishment can be… Species centred Resource centred …but in either case sufficient resources must be accumulated for an individual to establish a territory If territory establishment is… Species centred …then the ‘Position function” sets the parameters for territory establishment Territory establishment Saturation Half-saturation Territory densities may be… Low High …so realistic simulations must be calibrated to the real world To be as realistic as possible we had to make decisions concerning: •The characteristics of the landscape •The species’ distribution on thelandscape • The sampling method • The statistical model(s) Detection Function Point-count radius Vegetation plot radius To be as realistic as possible we had to make decisions concerning: •The characteristics of the landscape •The species’ distribution on thelandscape • The sampling method • The statistical model(s) The statistical model •Deterministic model structure Multiple regression, Logistic •Model error Normal, Poisson, Binomial •Model selection Parsimony (AIC), Bonferroni’s alpha, Statistical significance The deterministic model •Multiple regression (with 2 resources) Yi= B0 + B1X1i + B2X2i + B12X1iX2i + εi or Yi= f(X) + εi Yi = detection (0,1,2,…) X•i = resource value The deterministic model •Logarithmic: Yi= e f(X) + εi Yi = detection (0,1,2,...) X•i = resource value The deterministic model •Logistic: Yi= Ae f(X) /(1+ e f(X)) + εi Yi = detection (0,1,2,…) X•i = resource value Choosing the correct model form Linear model: 1 to 4 resources 1 Resource: Yi = B0 + B1X1i + εi 4 Resources: Yi = B0 + B1X1i + B2X2i + B3X3i + B4X4i Number of parameters required for… + B12X1iX2i + B13X1iX3i + B14X1iX4i 1 Resource = 2 + B23X2iX3i + B24X2iX4i + B34X3iX4i 2 Resource = 4 + B123X1iX2i X3i + B124X1iX2i X4i + B134X1iX3i X4i + B234X2iX3i X4i + B1234X1iX2i X3i X4i + εi 3 Resource = 8 4 Resource = 16 The statistical model •Deterministic model structure Multiple regression, Logistic •Model error Normal, Poisson, Binomial •Model selection Parsimony (AIC), Bonferroni’s alpha, Statistical significance Poisson error Repeated samples of individuals randomly dispersed are Poissondistributed Poisson error Negative-binomial error Normal error Binomial error The statistical model •Deterministic model structure Multiple regression, Logistic •Model error Normal, Poisson, Binomial •Model selection Parsimony (AIC), Bonferroni’s alpha, Statistical significance Model Selection •Use AIC to judge the best of several trial models •The ‘best’ model must be statistically significant from the ‘null’ model to be accepted If =0.05, then Bonferroni’s adjusted is: 1 Resource = 0.0500 2 Resource = .0169 3 Resource = 0.0073 4 Resource = 0.0034 True, Valid and Misleading Models •If the ‘True’ model is: Yi = B0 + B123X1iX2i X3i •Then: is a ‘Valid’ model •Yi = B0 + B3X3i •Yi = B0 + B12X1i X2i is a ‘Valid’ model •Yi = B0 + B4X4i •Yi = B0 + B14X1i X4i is a ‘Misleading’ model is a ‘Misleading’ model 1 Resource Required - 1 Resource Queried Success identifying ‘True’ Model Logistic-Poisson Multiple Regression - Normal 1 Resource Required - 1 Resource Queried Success identifying ‘True’ Model Logistic-Poisson Logistic-Binomial 4 Resources Required - 4 Resources Queried Medium SP - Resources uncorrelated – 100% detection - Full True Valid Misleading 4 Resources Required - 4 Resources Queried High SP - Resources uncorrelated – 100% detection - Full True Valid Misleading 4 Resources Required - 4 Resources Queried Low SP - Resources uncorrelated – 100% detection - Full True Valid Misleading 1 Resources Required - 4 Resources Queried Medium SP - Resources uncorrelated – 100% detection - Full True / Valid Misleading 1 Resources Required - 4 Resources Queried High SP - Resources uncorrelated – 100% detection - Full True / Valid Misleading 1 Resources Required - 4 Resources Queried Low SP - Resources uncorrelated – 100% detection - Full True / Valid Misleading 1 Resources Required - 4 Resources Queried Medium SP - Resources 50% correlated – 100% detection - Full True / Valid Misleading 1 Resources Required - 4 Resources Queried Medium SP - Resources 50% correlated – 25% detection - Full True / Valid Misleading 1 Resources Required - 4 Resources Queried Medium SP - Resources 50% correlated - 25% detection - 50% Full True / Valid Misleading 1 Resources Required - 4 Resources Queried High SP - Resources 50% correlated – 25% detection – 50% Full True / Valid Misleading 1 Resources Required - 4 Resources Queried Medium SP - Resources 95% correlated – 25% detection - Full True / Valid Misleading Technical Conclusions • A-priori hypotheses concerning species-habitat associations are essential • Required resources should be amalgamated by suite • Resource contrast is essential and should be planned: •Ratio of ‘between-point:within-point’ variability must be increased for both resources and species-of-interest •Point-count method must be designed with spatial period considerations in mind Key Conservation Conclusion At best: Affirmative conclusions about the importance of ‘critical resources’ based on statistical correlations alone are not justified! At worst: Affirmative conclusions about the importance of ‘critical resources’ based on statistical correlations alone, and without documenting the spatial characteristics of the landscape etc., are completely indefensible!