What we use is not what we know: environmental predictors in plant distribution models

Questions: The choice of environmental predictor variables in correlative models of plant species distributions (hereafter plant SDMs) is crucial to ensure predictive accuracy and model realism, as highlighted in multiple earlier studies. Because variable selection is directly related to a model's capacity to capture important species' environmental requirements, one would expect an explicit prior consideration of all ecophysiologically meaningful variables. For plants, these include temperature, water, soil nutrients, light, and in some cases, disturbances and biotic interactions. However, the set of predictors used in published correlative plant SDM studies varies considerably. No comprehensive review exists of what environmental predictors are meaningful, available (or missing), and used in practice to predict plant distributions. Contributing to answer these questions is the aim of this review. Methods: We carried out an extensive, systematic review of recently published plant SDM studies (years 2010-2015; n = 200) to determine the predictors used in the models. We additionally conducted an in-depth review of SDM studies in selected journals to identify temporal trends in the use of predictors (years 2000-2015; n = 40). Results: Except for the pure climatic studies, a large majority of plant SDM studies neglected several ecophysiologically-meaningful environmental variables, and the number of relevant predictors used in models has stagnated or even declined over the last 15 years. Conclusions: Neglecting ecophysiologically meaningful predictors can result in incomplete niche quantification and can thus limit the predictive power of plant SDMs. Some of these missing predictors are already available spatially or may soon become available (e.g., soil moisture). However, others are not yet easily obtainable across whole study extents (e.g., soil pH and nutrients), and their development should receive increased attention. We conclude that more effort should be made to build ecologically more sound plant SDMs. This requires a more thorough rationale for the choice of environmental predictors needed to meet the study goal, and the development of missing ones. The latter calls for increased collaborative effort between ecological and geo-environmental science

Climatic predictors of species distributions neglect biophysiologically meaningful variables

This is the final version. Available on open access from Wiley via the DOI in this record.Aim: Species distribution models (SDMs) have played a pivotal role in predicting how species might respond to climate change. To generate reliable and realistic predictions from these models requires the use of climate variables that adequately capture physiological responses of species to climate and therefore provide a proximal link between climate and their distributions. Here, we examine whether the climate variables used in plant SDMs are different from those known to influence directly plant physiology. Location: Global. Methods: We carry out an extensive, systematic review of the climate variables used to model the distributions of plant species and provide comparison to the climate variables identified as important in the plant physiology literature. We calculate the top ten SDM and physiology variables at 2.5 degree spatial resolution for the globe and use principal component analyses and multiple regression to assess similarity between the climatic variation described by both variable sets. Results: We find that the most commonly used SDM variables do not reflect the most important physiological variables and differ in two main ways: (i) SDM variables rely on seasonal or annual rainfall as simple proxies of water available to plants and neglect more direct measures such as soil water content; and (ii) SDM variables are typically averaged across seasons or years and overlook the importance of climatic events within the critical growth period of plants. We identify notable differences in their spatial gradients globally and show where distal variables may be less reliable proxies for the variables to which species are known to respond. Main conclusions: There is a growing need for the development of accessible, fine-resolution global climate surfaces of physiological variables. This would provide a means to improve the reliability of future range predictions from SDMs and support efforts to conserve biodiversity in a changing climate

A comprehensive evaluation of predictive performance of 33 species distribution models at species and community levels

A large array of species distribution model (SDM) approaches has been developed for explaining and predicting the occurrences of individual species or species assemblages. Given the wealth of existing models, it is unclear which models perform best for interpolation or extrapolation of existing data sets, particularly when one is concerned with species assemblages. We compared the predictive performance of 33 variants of 15 widely applied and recently emerged SDMs in the context of multispecies data, including both joint SDMs that model multiple species together, and stacked SDMs that model each species individually combining the predictions afterward. We offer a comprehensive evaluation of these SDM approaches by examining their performance in predicting withheld empirical validation data of different sizes representing five different taxonomic groups, and for prediction tasks related to both interpolation and extrapolation. We measure predictive performance by 12 measures of accuracy, discrimination power, calibration, and precision of predictions, for the biological levels of species occurrence, species richness, and community composition. Our results show large variation among the models in their predictive performance, especially for communities comprising many species that are rare. The results do not reveal any major trade-offs among measures of model performance; the same models performed generally well in terms of accuracy, discrimination, and calibration, and for the biological levels of individual species, species richness, and community composition. In contrast, the models that gave the most precise predictions were not well calibrated, suggesting that poorly performing models can make overconfident predictions. However, none of the models performed well for all prediction tasks. As a general strategy, we therefore propose that researchers fit a small set of models showing complementary performance, and then apply a cross-validation procedure involving separate data to establish which of these models performs best for the goal of the study.Peer reviewe

Biotic interactions boost spatial models of species richness

Biotic interactions are known to affect the composition of species assemblages via several mechanisms, such as competition and facilitation. However, most spatial models of species richness do not explicitly consider inter-specific interactions. Here, we test whether incorporating biotic interactions into high-resolution models alters predictions of species richness as hypothesised. We included key biotic variables (cover of three dominant arctic-alpine plant species) into two methodologically divergent species richness modelling frameworks - stacked species distribution models (SSDM) and macroecological models (MEM) - for three ecologically and evolutionary distinct taxonomic groups (vascular plants, bryophytes and lichens). Predictions from models including biotic interactions were compared to the predictions of models based on climatic and abiotic data only. Including plant-plant interactions consistently and significantly lowered bias in species richness predictions and increased predictive power for independent evaluation data when compared to the conventional climatic and abiotic data based models. Improvements in predictions were constant irrespective of the modelling framework or taxonomic group used. The global biodiversity crisis necessitates accurate predictions of how changes in biotic and abiotic conditions will potentially affect species richness patterns. Here, we demonstrate that models of the spatial distribution of species richness can be improved by incorporating biotic interactions, and thus that these key predictor factors must be accounted for in biodiversity forecast

A comprehensive evaluation of predictive performance of 33 species distribution models at species and community levels

International audienc