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Predictive performance of presence‐only species distribution models: a benchmark study with reproducible code

Valavi, Roozbeh, Guillera‐Arroita, Gurutzeta, Lahoz‐Monfort, José J., Elith, Jane·2022·Ecological Monographs

Abstract

Species distribution modeling (SDM) is widely used in ecology and conservation. Currently, the most available data for SDM are species presence‐only records (available through digital databases). There have been many studies comparing the performance of alternative algorithms for modeling presence‐only data. Among these, a 2006 paper from Elith and colleagues has been particularly influential in the field, partly because they used several novel methods (at the time) on a global data set that included independent presence–absence records for model evaluation. Since its publication, some of the algorithms have been further developed and new ones have emerged. In this paper, we explore patterns in predictive performance across methods, by reanalyzing the same data set (225 species from six different regions) using updated modeling knowledge and practices. We apply well‐established methods such as generalized additive models and MaxEnt, alongside others that have received attention more recently, including regularized regressions, point‐process weighted regressions, random forests, XGBoost, support vector machines, and the ensemble modeling framework biomod. All the methods we use include background samples (a sample of environments in the landscape) for model fitting. We explore impacts of using weights on the presence and background points in model fitting. We introduce new ways of evaluating models fitted to these data, using the area under the precision‐recall gain curve, and focusing on the rank of results. We find that the way models are fitted matters. The top method was an ensemble of tuned individual models. In contrast, ensembles built using the biomod framework with default parameters performed no better than single moderate performing models. Similarly, the second top performing method was a random forest parameterized to deal with many background samples (contrasted to relatively few presence records), which substantially outperformed other random forest implementations. We find that, in general, nonparametric techniques with the capability of controlling for model complexity outperformed traditional regression methods, with MaxEnt and boosted regression trees still among the top performing models. All the data and code with working examples are provided to make this study fully reproducible.

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Image 1
CC-BY-NC-ND · From the PDF
FIG. 2 AUCROC values for all modeling methods across all species. Each circular track is the AUCROC of a model for all species. Models and species are ordered by average AUCROC. The outer text labels indicate the regions. The numbers between NSW and NZ regions label the modeling method as mentioned in middle of the figure. The height of the outer histogram shows the log10(number of species presence-only records) in the training data set. Histogram colors follow three categories: light orange. < 100; pink, 100 -1,000; and dark brown, > 1,000 presences. This figure was created in Circos software (Krzywinski et al. 2009).
FIG. 2 AUCROC values for all modeling methods across all species. Each circular track is the AUCROC of a model for all species. Models and species are ordered by average AUCROC. The outer text labels indicate the regions. The numbers between NSW and NZ regions label the modeling method as mentioned in middle of the figure. The height of the outer histogram shows the log10(number of species presence-only records) in the training data set. Histogram colors follow three categories: light orange. < 100; pink, 100 -1,000; and dark brown, > 1,000 presences. This figure was created in Circos software (Krzywinski et al. 2009).CC-BY-NC-ND · From the PDF
FIG. 3 Mean AUCROC vs. mean COR for the models, summarized across all species. The gray bars (around each model) are standard errors estimated in a Bayesian mixed model, reflecting variation for an average species in an average region. The solid gray line is the least-squares line fitted on the models. The dashed gray lines delimit regions of low-, moderate-, and high-performance models, following the hierarchical clustering (represented in the top of the plot in Fig. 4). As a reminder, the ensemble model is the average of Lasso, GAM, MaxEnt, BRT, and RF down-sampled.
FIG. 3 Mean AUCROC vs. mean COR for the models, summarized across all species. The gray bars (around each model) are standard errors estimated in a Bayesian mixed model, reflecting variation for an average species in an average region. The solid gray line is the least-squares line fitted on the models. The dashed gray lines delimit regions of low-, moderate-, and high-performance models, following the hierarchical clustering (represented in the top of the plot in Fig. 4). As a reminder, the ensemble model is the average of Lasso, GAM, MaxEnt, BRT, and RF down-sampled.CC-BY-NC-ND · From the PDF
FIG. 4 P values of the post-hoc of the Friedman Aligned Rank test with Shaffer adjustments for AUCROC for a null hypothesis of no difference between pairs of models (values are rounded). The colors and the text inside each cell show different P values. Darker colors indicate higher P values (no significant difference in performance between models). The dendrogram on the top is based on both AUCROC and COR that clusters models in three main groups. The height of each bar shows the relative difference between each model/group.
FIG. 4 P values of the post-hoc of the Friedman Aligned Rank test with Shaffer adjustments for AUCROC for a null hypothesis of no difference between pairs of models (values are rounded). The colors and the text inside each cell show different P values. Darker colors indicate higher P values (no significant difference in performance between models). The dendrogram on the top is based on both AUCROC and COR that clusters models in three main groups. The height of each bar shows the relative difference between each model/group.CC-BY-NC-ND · From the PDF
FIG. 5 The difference between the performance of the default model with modified models in RF, SVM, and ensemble models. Other methods are presented in gray for reference. Bars and lines as in Fig. 3.
FIG. 5 The difference between the performance of the default model with modified models in RF, SVM, and ensemble models. Other methods are presented in gray for reference. Bars and lines as in Fig. 3.CC-BY-NC-ND · From the PDF
FIG. 6 Performance of models in NCEAS 2006, Phillips et al. (2017), and this study. For ease of comparison, this plot uses the same axes ranges and division lines as Fig. 3. The dashed and solid gray lines separate the low-, moderate-, and high-performance models, as in Fig. 3. The Phillips et al. (2017) models have no standard errors attached, as only their average performance was available to us. The MAXENTT from NCEAS 2006 study is a MaxEnt model with ' threshold ' feature (this feature is not included in the other studies).
FIG. 6 Performance of models in NCEAS 2006, Phillips et al. (2017), and this study. For ease of comparison, this plot uses the same axes ranges and division lines as Fig. 3. The dashed and solid gray lines separate the low-, moderate-, and high-performance models, as in Fig. 3. The Phillips et al. (2017) models have no standard errors attached, as only their average performance was available to us. The MAXENTT from NCEAS 2006 study is a MaxEnt model with ' threshold ' feature (this feature is not included in the other studies).CC-BY-NC-ND · From the PDF
FIG. 7 The average rank of AUCROC vs. (A) Pearson correlation between the predicted likelihood of presence and the presence -absence testing data (COR) and (B) area under the precision-recall gain curve (AUCPRG) for all the models. The gray dotted lines are the mean ranks. The values of the x and y -axis are reversed to have the lowest rank (better models) on the top-right corner and vice versa.
FIG. 7 The average rank of AUCROC vs. (A) Pearson correlation between the predicted likelihood of presence and the presence -absence testing data (COR) and (B) area under the precision-recall gain curve (AUCPRG) for all the models. The gray dotted lines are the mean ranks. The values of the x and y -axis are reversed to have the lowest rank (better models) on the top-right corner and vice versa.CC-BY-NC-ND · From the PDF
FIG. 9 Mean and standard error of the AUCROC for the species with < 30 and ≥ 30 presence points (with 61 and 164 species, respectively). Models are arranged based on median AUCROC in their corresponding group: tree-based, regression, and others.
FIG. 9 Mean and standard error of the AUCROC for the species with < 30 and ≥ 30 presence points (with 61 and 164 species, respectively). Models are arranged based on median AUCROC in their corresponding group: tree-based, regression, and others.CC-BY-NC-ND · From the PDF
Image 9
CC-BY-NC-ND · From the PDF
FIG. 11 Map of predicted relative likelihood of occurrence for the species nz30 in the NZ data set. The top-left figure shows the occurrence data used for fitting the models. The colored histogram to the left of each map shows the distribution of predicted values on all grid cells and the transparent gray histogram on top of it, demonstrates the distribution of predicted values on presence points (from testing data). The SVM weighted predicted values were very low, so its map is linearly rescaled between 0 and 1.
FIG. 11 Map of predicted relative likelihood of occurrence for the species nz30 in the NZ data set. The top-left figure shows the occurrence data used for fitting the models. The colored histogram to the left of each map shows the distribution of predicted values on all grid cells and the transparent gray histogram on top of it, demonstrates the distribution of predicted values on presence points (from testing data). The SVM weighted predicted values were very low, so its map is linearly rescaled between 0 and 1.CC-BY-NC-ND · From the PDF
FIG. 12 Model-fitting computation time for each method. Methods are arranged along the x -axis based on the median of times (from short to long). The ensemble model runtime was calculated by summing the computation times for all its underlying models (see Modeling methods ). XGBoost and MARS models were run with multiple CPU cores in parallel (eight cores).
FIG. 12 Model-fitting computation time for each method. Methods are arranged along the x -axis based on the median of times (from short to long). The ensemble model runtime was calculated by summing the computation times for all its underlying models (see Modeling methods ). XGBoost and MARS models were run with multiple CPU cores in parallel (eight cores).CC-BY-NC-ND · From the PDF

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Metadata

Authors
Valavi, R., Guillera‐Arroita, G., Lahoz‐Monfort, J. J., & Elith, J.
Venue
Ecological Monographs
Publisher
Wiley
Published
2021-11-16
Volume
92
Issue
1
ISSN
  • 0012-9615
  • 1557-7015
Cited by
830
References
6
Language
en

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