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ECOLOGICAL NICHE MODELING METHODS
UPDATE
Town Peterson, University of Kansas
It Is A Bit Too Easy …
• Very easy access to lots of
occurrence data
• Very easy access to rich
geospatial data
• Easy-to-use modeling tools
• Lots of literature setting out
the examples
Ecological Niche Modeling
1. Accumulate Input Data
2. Integrate Occurrence and Environmental Data
3. Model Calibration
4. Model Evaluation
5. Summary and Interpretation
Accumulate Input Data
Collate primary
biodiversity data
documenting
occurrences
Process environmental
layers to be maximally
relevant to distributional
ecology of species in
question
Collate GIS database
of relevant data
layers
Assess spatial precision
of occurrence data;
adjust inclusion of data
accordingly
Data subsetting for
model evaluation
Occurrence and
environmental data
Assess spatial
autocorrelation
Occurrence Data in Niche Modeling
• Goal is to represent the full diversity of
situations under which a particular species
maintains populations
• Spatial biases (i.e., non-random or non-
uniform distribution within G) is not damning
• Biases within E are catastrophic, and will
translate directly into biases in any niche
estimate
• More is usually better, but not always…
speciesLink Network
Uncertainty in Direction and Distance
Georeferencing should …
• Represent the place at which the species was
found
• Represent the certainty and uncertainty with
which that place is characterized
• Summarize the methods used to establish that
place
• Preserve all of the original information for
possible reinterpretation
Internal Consistency Testing
Data Cleaning
• Attempt to detect meaningfully erroneous records,
so that they can be treated with caution in analysis
• Use internal consistency to detect initial problems
– Species names consistent?
– Terrestrial species on land, marine species in the ocean?
– Latlong matches country, state, district, etc.
• Use external consistency to go deeper
– Occurrence data match known distribution spatially?
– Occurrence data match known distribution
environmentally?
• If precision data are available, filter to retain only
records that are precise enough for the study
• Iterative process with important consequences
Data Subsetting
• Must respond to the question at hand … why
are you doing the study?
• Ideally completely independent data streams
• Failing that, can be
– Macrospatial
– Microspatial (but see spatial autocorrelation)
– Random
• Will return to this point later…
Generalities: Environmental Data
• Raster format: i.e., information exists across
entire region of interest
• Relevant information as regards the
distributional potential of the species of
interest
• More dimensions = better (generally), BUT
– collinearity is bad
– too many dimensions is bad
Major Sources
• Climate data – long time span, but low
temporal resolution
• Remote-sensing data – high temporal
resolution, diverse products, short time span
• Topographic data – high temporal resolution,
uncertain connection to species’ distributional
ecology
• Soils data – uneven global coverage,
categorical data
• Others
Spatial
Autocorrelation
Two Major Implications
• Non-independence in model evaluation
– Available data are often split into data sets for
calibration and evaluation
– Data points that are not independent of one another
may end up in different data sets, thereby
compromising the robustness of the test
• Inflation of sample sizes
– Because individual data points may be non-
independent of one another, sample sizes may appear
larger than they actually are
– This inflation may create opportunity for Type 1 errors
in model evaluation and model comparisons
Process for Maximum Relevancy
Integrate Occurrence and Environmental Data
Assess BAM scenario for
species in question; avoid
M-limited situations
Saupe et al. 2012. Variation in niche and distribution model performance: The need
for a priori assessment of key causal factors. Ecological Modelling, 237–238, 11-22.
Estimate M and S
as area of analysis
in study
Barve et al. 2011. The crucial role of the
accessible area in ecological niche modeling
and species distribution modeling. Ecological
Modelling, 222, 1810-1819.
Reduce dimensionality (PCA
or correlation analysis)
Occurrence and
environmental data
Occurrence and
environmental data
ready for analysis
Assess BAM Scenario
BAM I: Eltonian Noise Hypothesis
A
M
B A
M
BAM II
Classic
BAM
Hutchinson’s
Dream
Wallace’s
Dream
All OK
Project onto Geography
Effect of BAM Scenarios
-0.1
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
-0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1
-0.1
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
-0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1
-0.1
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
-0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1
CB
EF
WD
-0.1
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
-0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1
HD
KGo
KAKA
KGo
KGoKGo
KA KA
Figure 3. CB = Classic BAM scenario; WD = Wallace’s Dream; EF = Elith’s Fantasy; HD = Hutchinson’s Dream
Blue = Virtual Species 1; Green = Virtual Species 3; Orange = Virtual Species 4
Circles = GARP; Squares = Maxent; Triangles = GAM
BAM Conclusions
• Some situations are not amenable to fitting
ecological niche models that will have
predictive power
• Models tend much more to good fitting of the
potential distribution, rather than the actual
distribution
• Must ponder carefully the BAM configuration
in a particular study situation to avoid
configurations that will not yield usable
models
M and S as Study Area
Test Arena: The Lawrence Species
M and Model Training
Model Evaluation
M and Model Comparison
Model Comparison
M
• When the species has no history in an area:
– Use a radius related to dispersal distances
• When history is short (i.e., environment
constant):
– Use a radius representing compounding of
dispersal distances
• When history is long (i.e., environmental
change is a factor)
– Seek ways of assessing areas that the species’
distribution through time has covered…
Icterus cucullatus
Sampling
Reduce Dimensionality
Model Calibration
Estimate ecological niche
(various algorithms)
Model calibration,
adjusting parameters to
maximize quality
Model
thresholding
Peterson et al. 2007. Transferability and model
evaluation in ecological niche modeling: A
comparison of GARP and Maxent. Ecography,
30, 550-560.
Occurrence and
environmental data
ready for analysis
“No Silver Bullet” paper to appear
Warren, D. L. and S. N. Seifert. 2011. Ecological niche
modeling in Maxent: The importance of model complexity
and the performance of model selection criteria. Ecological
Applications 21:335-342.
Preliminary
models
Estimate Ecological Niche
No Silver Bullets in ENM
• Single algorithms may perform ‘best’ on average
• The best algorithm in any given situation,
however, may be other than the ‘best’
• NSB thinking suggests that we should not use a
single approach
• Use a suite of approaches (e.g., as implemented
in OM, BIOMOD, BIOENSEMBLES, etc.), challenge
to predict, choose best for that situation
• Maxent is good, but it is not the only algorithm …
Model Thresholding
“Presence”
Thresholding
• Use an approach that prioritizes omission
error over commission error, in view of the
greater reliability of presence data
• Minimum training presence thresholding
seeks the highest suitability value that
includes 100% of the calibration data
• Suggest (strongly) using a parallel approach
that seeks that highest suitability value that
includes (100-E)% of the calibration data
Model Optimization and Parameter Choice
Model Evaluation
Project niche model
to geographic space
Model
evaluation
Peterson et al. 2008. Rethinking receiver operating characteristic
analysis applications in ecological niche modelling. Ecological
Modelling, 213, 63-72.
Preliminary
models
Reset data subsets based
on evaluation results
Corroborated models
ready for projection to
geographic
times/regions of
interest
If predicted suitable area covers
15% of the testing area, then
15% of evaluation points are
expected to fall in the predicted
suitable area by chance.
• p = proportion of area
predicted suitable
• s = number of successes
• n = number of evaluation
points
Cumulative binomial distribution calculates the probability of
obtaining s successes out of n trials in a situation in which p
proportion of the testing area is predicted present. If this probability
is below 0.05, we interpret the situation as indicating that the
model’s predictions are significantly better than random.
Threshold-
dependent
Approach
Threshold-independent Approaches
http://shiny.conabio.gob.mx:3838/nichetoolb2/
Significance vs Performance
• Predictions that are significantly better than
random is important, and is a sine qua non for
model interpretation
• BUT, it is also important to assure that the
model performs sufficiently well for the
intended uses of the output
• Performance measures include omission rate,
correct classification rate, etc.
Summary and Interpretation
Evaluation of model
transfer results
Transfer to other
situations (time
and space)
Assess extrapolation
(MESS and MOP)
Owens, H. L., L. P. Campbell, L. Dornak, E. E. Saupe, N.
Barve, J. Soberón, K. Ingenloff, A. Lira-Noriega, C. M.
Hensz, C. E. Myers, and A. T. Peterson. 2013.
Constraints on interpretation of ecological niche
models by limited environmental ranges on calibration
areas. Ecological Modelling 263:10-18.
Refine estimate of
current distribution
via land use, etc.
Compare present and
“other” to assess
effects of change
Models calibrated and
evaluated, and transferred
to present and “other”
situations
MESS and MOP
• Both have the intention of detecting extrapolative
situations
• MESS is implemented within Maxent
• MESS compares the area in question to the
centroid of the calibration cloud
• MOP compares the area in question to the
nearest part of the calibration cloud
• Agree on ‘out of range’ conditions
• MOP better characterizes similarities between
calibration and transfer regions, and thus is more
optimistic as regards in-range extrapolation
Ecological Niche Modeling
1. Accumulate Input Data
2. Integrate Occurrence and Environmental Data
3. Model Calibration
4. Model Evaluation
5. Summary and Interpretation
town@ku.edu

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Updating Ecological Niche Modeling Methodologies

  • 1. ECOLOGICAL NICHE MODELING METHODS UPDATE Town Peterson, University of Kansas
  • 2. It Is A Bit Too Easy … • Very easy access to lots of occurrence data • Very easy access to rich geospatial data • Easy-to-use modeling tools • Lots of literature setting out the examples
  • 3. Ecological Niche Modeling 1. Accumulate Input Data 2. Integrate Occurrence and Environmental Data 3. Model Calibration 4. Model Evaluation 5. Summary and Interpretation
  • 4. Accumulate Input Data Collate primary biodiversity data documenting occurrences Process environmental layers to be maximally relevant to distributional ecology of species in question Collate GIS database of relevant data layers Assess spatial precision of occurrence data; adjust inclusion of data accordingly Data subsetting for model evaluation Occurrence and environmental data Assess spatial autocorrelation
  • 5. Occurrence Data in Niche Modeling • Goal is to represent the full diversity of situations under which a particular species maintains populations • Spatial biases (i.e., non-random or non- uniform distribution within G) is not damning • Biases within E are catastrophic, and will translate directly into biases in any niche estimate • More is usually better, but not always…
  • 8. Georeferencing should … • Represent the place at which the species was found • Represent the certainty and uncertainty with which that place is characterized • Summarize the methods used to establish that place • Preserve all of the original information for possible reinterpretation
  • 9.
  • 10.
  • 12. Data Cleaning • Attempt to detect meaningfully erroneous records, so that they can be treated with caution in analysis • Use internal consistency to detect initial problems – Species names consistent? – Terrestrial species on land, marine species in the ocean? – Latlong matches country, state, district, etc. • Use external consistency to go deeper – Occurrence data match known distribution spatially? – Occurrence data match known distribution environmentally? • If precision data are available, filter to retain only records that are precise enough for the study • Iterative process with important consequences
  • 13.
  • 14. Data Subsetting • Must respond to the question at hand … why are you doing the study? • Ideally completely independent data streams • Failing that, can be – Macrospatial – Microspatial (but see spatial autocorrelation) – Random • Will return to this point later…
  • 15. Generalities: Environmental Data • Raster format: i.e., information exists across entire region of interest • Relevant information as regards the distributional potential of the species of interest • More dimensions = better (generally), BUT – collinearity is bad – too many dimensions is bad
  • 16. Major Sources • Climate data – long time span, but low temporal resolution • Remote-sensing data – high temporal resolution, diverse products, short time span • Topographic data – high temporal resolution, uncertain connection to species’ distributional ecology • Soils data – uneven global coverage, categorical data • Others
  • 18. Two Major Implications • Non-independence in model evaluation – Available data are often split into data sets for calibration and evaluation – Data points that are not independent of one another may end up in different data sets, thereby compromising the robustness of the test • Inflation of sample sizes – Because individual data points may be non- independent of one another, sample sizes may appear larger than they actually are – This inflation may create opportunity for Type 1 errors in model evaluation and model comparisons
  • 19.
  • 20. Process for Maximum Relevancy
  • 21. Integrate Occurrence and Environmental Data Assess BAM scenario for species in question; avoid M-limited situations Saupe et al. 2012. Variation in niche and distribution model performance: The need for a priori assessment of key causal factors. Ecological Modelling, 237–238, 11-22. Estimate M and S as area of analysis in study Barve et al. 2011. The crucial role of the accessible area in ecological niche modeling and species distribution modeling. Ecological Modelling, 222, 1810-1819. Reduce dimensionality (PCA or correlation analysis) Occurrence and environmental data Occurrence and environmental data ready for analysis
  • 23. BAM I: Eltonian Noise Hypothesis A M B A M
  • 26. Effect of BAM Scenarios -0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 -0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 -0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 -0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 -0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 -0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 CB EF WD -0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 -0.1 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1 HD KGo KAKA KGo KGoKGo KA KA Figure 3. CB = Classic BAM scenario; WD = Wallace’s Dream; EF = Elith’s Fantasy; HD = Hutchinson’s Dream Blue = Virtual Species 1; Green = Virtual Species 3; Orange = Virtual Species 4 Circles = GARP; Squares = Maxent; Triangles = GAM
  • 27. BAM Conclusions • Some situations are not amenable to fitting ecological niche models that will have predictive power • Models tend much more to good fitting of the potential distribution, rather than the actual distribution • Must ponder carefully the BAM configuration in a particular study situation to avoid configurations that will not yield usable models
  • 28. M and S as Study Area
  • 29. Test Arena: The Lawrence Species
  • 30. M and Model Training
  • 32. M and Model Comparison
  • 34. M • When the species has no history in an area: – Use a radius related to dispersal distances • When history is short (i.e., environment constant): – Use a radius representing compounding of dispersal distances • When history is long (i.e., environmental change is a factor) – Seek ways of assessing areas that the species’ distribution through time has covered…
  • 37. Model Calibration Estimate ecological niche (various algorithms) Model calibration, adjusting parameters to maximize quality Model thresholding Peterson et al. 2007. Transferability and model evaluation in ecological niche modeling: A comparison of GARP and Maxent. Ecography, 30, 550-560. Occurrence and environmental data ready for analysis “No Silver Bullet” paper to appear Warren, D. L. and S. N. Seifert. 2011. Ecological niche modeling in Maxent: The importance of model complexity and the performance of model selection criteria. Ecological Applications 21:335-342. Preliminary models
  • 39.
  • 40.
  • 41.
  • 42.
  • 43.
  • 44.
  • 45.
  • 46.
  • 47. No Silver Bullets in ENM • Single algorithms may perform ‘best’ on average • The best algorithm in any given situation, however, may be other than the ‘best’ • NSB thinking suggests that we should not use a single approach • Use a suite of approaches (e.g., as implemented in OM, BIOMOD, BIOENSEMBLES, etc.), challenge to predict, choose best for that situation • Maxent is good, but it is not the only algorithm …
  • 50. Thresholding • Use an approach that prioritizes omission error over commission error, in view of the greater reliability of presence data • Minimum training presence thresholding seeks the highest suitability value that includes 100% of the calibration data • Suggest (strongly) using a parallel approach that seeks that highest suitability value that includes (100-E)% of the calibration data
  • 51. Model Optimization and Parameter Choice
  • 52. Model Evaluation Project niche model to geographic space Model evaluation Peterson et al. 2008. Rethinking receiver operating characteristic analysis applications in ecological niche modelling. Ecological Modelling, 213, 63-72. Preliminary models Reset data subsets based on evaluation results Corroborated models ready for projection to geographic times/regions of interest
  • 53.
  • 54. If predicted suitable area covers 15% of the testing area, then 15% of evaluation points are expected to fall in the predicted suitable area by chance. • p = proportion of area predicted suitable • s = number of successes • n = number of evaluation points Cumulative binomial distribution calculates the probability of obtaining s successes out of n trials in a situation in which p proportion of the testing area is predicted present. If this probability is below 0.05, we interpret the situation as indicating that the model’s predictions are significantly better than random. Threshold- dependent Approach
  • 56.
  • 57.
  • 59. Significance vs Performance • Predictions that are significantly better than random is important, and is a sine qua non for model interpretation • BUT, it is also important to assure that the model performs sufficiently well for the intended uses of the output • Performance measures include omission rate, correct classification rate, etc.
  • 60. Summary and Interpretation Evaluation of model transfer results Transfer to other situations (time and space) Assess extrapolation (MESS and MOP) Owens, H. L., L. P. Campbell, L. Dornak, E. E. Saupe, N. Barve, J. Soberón, K. Ingenloff, A. Lira-Noriega, C. M. Hensz, C. E. Myers, and A. T. Peterson. 2013. Constraints on interpretation of ecological niche models by limited environmental ranges on calibration areas. Ecological Modelling 263:10-18. Refine estimate of current distribution via land use, etc. Compare present and “other” to assess effects of change Models calibrated and evaluated, and transferred to present and “other” situations
  • 61.
  • 62.
  • 63.
  • 64.
  • 65.
  • 66. MESS and MOP • Both have the intention of detecting extrapolative situations • MESS is implemented within Maxent • MESS compares the area in question to the centroid of the calibration cloud • MOP compares the area in question to the nearest part of the calibration cloud • Agree on ‘out of range’ conditions • MOP better characterizes similarities between calibration and transfer regions, and thus is more optimistic as regards in-range extrapolation
  • 67.
  • 68. Ecological Niche Modeling 1. Accumulate Input Data 2. Integrate Occurrence and Environmental Data 3. Model Calibration 4. Model Evaluation 5. Summary and Interpretation