Accounting for multi‐scale spatial autocorrelation improves performance of invasive species distribution modelling (iSDM)

Aim Analyses of species distributions are complicated by various origins of spatial autocorrelation (SAC) in biogeographical data. SAC may be particularly important for invasive species distribution models (iSDMs) because biological invasions are strongly influenced by dispersal and colonization processes that typically create highly structured distribution patterns. We examined the efficacy of using a multi‐scale framework to account for different origins of SAC, and compared non‐spatial models with models that accounted for SAC at multiple levels. Location We modelled the spatial distribution of an invasive forest pathogen, Phytophthora ramorum, in western USA. Methods We applied one conventional statistical method (generalized linear model, GLM) and one nonparametric technique (maximum entropy, Maxent) to a large dataset on P. ramorum occurrence (n = 3787) to develop four types of model that included environmental variables and that either ignored spatial context or incorporated it at a broad scale using trend surface analysis, a local scale using autocovariates, or multiple scales using spatial eigenvector mapping. We evaluated model accuracies and amounts of explained spatial structure, and examined the changes in predictive power of the environmental and spatial variables. Results Accounting for different scales of SAC significantly enhanced the predictive capability of iSDMs. Dramatic improvements were observed when fine‐scale SAC was included, suggesting that local range‐confining processes are important in P. ramorum spread. The importance of environmental variables was relatively consistent across all models, but the explanatory power decreased in spatial models for factors with strong spatial structure. While accounting for SAC reduced the amount of residual autocorrelation for GLM but not for Maxent, it still improved the performance of both approaches, supporting our hypothesis that dispersal and colonization processes are important factors to consider in distribution models of biological invasions. Main conclusions Spatial autocorrelation has become a paradigm in biogeography and ecological modelling. In addition to avoiding the violation of statistical assumptions, accounting for spatial patterns at multiple scales can enhance our understanding of dynamic processes that explain ecological mechanisms of invasion and improve the predictive performance of static iSDMs.     

东北地区5个物种潜在栖息地变化与优化保护规划

气候变化广泛影响着物种多样性及其分布变迁。优化模型模拟结果,获取气候变化影响下的优先保护区域将为制定应对气候变化的物种保护政策或行动提供理论依据,提升保护绩效。选取东北地区五种代表性动物,包括黑熊(Ursus thibetanus)、驼鹿(Alces alces)、水獭(Lutra lutra)、紫貂(Martes zibellina)及黑嘴松鸡(Tetrao parvirostris);结合最大熵模型(Maxent)模拟在不同RCP情景下未来3个年代(2030s,2050s,2070s)的物种潜在栖息地。根据九个常用气候模式的评价结果,获取东北地区合适的气候模式,了解气候变化对物种潜在栖息地的影响,同时开展物种保护规划,识别保护空缺,为应对气候变化、保持生物多样性提供支持。结果显示,在气候变化背景下物种潜在栖息地面积整体呈现下降趋势,但不同气候模式之间存在差异;评价结果推荐CCSM4、Nor ESM1-M、Had GEM2-AO及GFDL-CM3气候模式,推荐在东北地区使用以上气候模式进行物种未来潜在分布的研究。5个物种潜在栖息地平均面积变化率分别为-62.16%,-73.93%,-78.46%(2030s,2050s,2070s)。综合5个重点保护物种的保护优先区,大兴安岭的呼中、汗马与额尔古纳国家级自然保护区,延边地区的天佛指山、老爷岭东北虎、珲春东北虎与汪清原麝国家级自然保护区,长白山国家级自然保护区是气候变化下物种保护的热点区域。     

Spatial autocorrelation in predictors reduces the impact of positional uncertainty in occurrence data on species distribution modelling

Aim To investigate the impact of positional uncertainty in species occurrences on the predictions of seven commonly used species distribution models (SDMs), and explore its interaction with spatial autocorrelation in predictors. Methods A series of artificial datasets covering 155 scenarios including different combinations of five positional uncertainty scenarios and 31 spatial autocorrelation scenarios were simulated. The level of positional uncertainty was defined by the standard deviation of a normally distributed zero‐mean random variable. Each dataset included two environmental gradients (predictor variables) and one set of species occurrence sample points (response variable). Seven commonly used models were selected to develop SDMs: generalized linear models, generalized additive models, boosted regression trees, multivariate adaptive regression spline, random forests, genetic algorithm for rule‐set production and maximum entropy. A probabilistic approach was employed to model and simulate five levels of error in the species locations. To analyse the propagation of positional uncertainty, Monte Carlo simulation was applied to each scenario for each SDM. The models were evaluated for performance using simulated independent test data with Cohen’s Kappa and the area under the receiver operating characteristic curve. Results Positional uncertainty in species location led to a reduction in prediction accuracy for all SDMs, although the magnitude of the reduction varied between SDMs. In all cases the magnitude of this impact varied according to the degree of spatial autocorrelation in predictors and the levels of positional uncertainty. It was shown that when the range of spatial autocorrelation in the predictors was less than or equal to three times the standard deviation of the positional error, the models were less affected by error and, consequently, had smaller decreases in prediction accuracy. When the range of spatial autocorrelation in predictors was larger than three times the standard deviation of positional error, the prediction accuracy was low for all scenarios. Main conclusions The potential impact of positional uncertainty in species occurrences on the predictions of SDMs can be understood by comparing it with the spatial autocorrelation range in predictor variables.     

The performance of range maps and species distribution models representing the geographic variation of species richness at different resolutions

Aim The method used to generate hypotheses about species distributions, in addition to spatial scale, may affect the biodiversity patterns that are then observed. We compared the performance of range maps and MaxEnt species distribution models at different spatial resolutions by examining the degree of similarity between predicted species richness and composition against observed values from well‐surveyed cells (WSCs). Location Mexico. Methods We estimated amphibian richness distributions at five spatial resolutions (from 0.083° to 2°) by overlaying 370 individual range maps or MaxEnt predictions, comparing the similarity of the spatial patterns and correlating predicted values with the observed values for WSCs. Additionally, we looked at species composition and assessed commission and omission errors associated with each method. Results MaxEnt predictions reveal greater geographic differences in richness between species rich and species poor regions than the range maps did at the five resolutions assessed. Correlations between species richness values estimated by either of the two procedures and the observed values from the WSCs increased with decreasing resolution. The slopes of the regressions between the predicted and observed values indicate that MaxEnt overpredicts observed species richness at all of the resolutions used, while range maps underpredict them, except at the finest resolution. Prediction errors did not vary significantly between methods at any resolution and tended to decrease with decreasing resolution. The accuracy of both procedures was clearly different when commission and omission errors were examined separately. Main conclusions Despite the congruent increase in the geographic richness patterns obtained from both procedures as resolution decreases, the maps created with these methods cannot be used interchangeably because of notable differences in the species compositions they report.     

中国植物分布模拟研究现状

在过去的20年里, 物种分布模型已广泛应用于动植物地理分布的模拟研究。该文以植物物种分布模拟为例, 利用中国知网、维普网以及Web of Science文献数据库的检索与统计, 分析了2000-2018年间, 中国研究人员利用各种物种分布模型对植物物种分布模拟研究的发文量、模拟模型、物种类型、数据来源、研究目的等信息。最终共收集到366篇有效文献, 分析表明2011年以来中国的物种分布模型应用发展迅速, 且以最近5年最为迅猛, 在生态学、中草药业、农业和林业等行业部门应用广泛。在使用的33种模型中, 应用最广的为最大熵模型(MaxEnt)。有一半研究的环境数据仅包含气候数据, 另一半研究不仅包含气候数据还包括地形与土壤等数据; 环境及物种数据的来源多样, 国际及本土数据库均得到使用。模拟涉及有明确清单的562个植物种, 既有木本植物(52.7%), 也有草本植物(41.8%), 其中中草药、果树、园林植物、农作物等占比较高。研究目的主要集中在过去、现在和未来气候变化对植物种分布的影响及预测, 以及物种分布评估与生物多样性评价(包括入侵植物风险评估)两大方面。预测物种潜在分布范围与气候变化影响等基础研究, 与模拟物种适生区与推广种植等应用研究并重, 物种分布模型在生态学与农业、林业和中草药业等多学科、多行业开展多种应用, 多物种、多模型和多来源数据共同参与模拟与比较, 开发新的机理性物种分布模型, 拓展新的物种分布模拟应用领域, 是今后研究的重点发展方向。     

Geographical distribution modelling of the bronze bug: a worldwide invasion

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Climate controls the distribution of a widespread invasive species: implications for future range expansion

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Habitat distribution modelling,under the present climatic scenario,of the threatened endemic Iberian Delphinium fissum subsp. sordidum (Ranunculaceae) and implications for its conservation

We modelled the potential habitat of a threatened species D. fissum subsp. sordidum, an endemic hemicryptophyte with a disjunct distribution in the Iberian Peninsula. Maxent was used to predict the subspecies habitat suitability by relating field sample-based distributional information with environmental and topographic variables. Our results suggest that the model performed well, predicting with high accuracy the current distribution of the species. The variables that most contributed to the model were Mean Temperature of Wettest Quarter (MTWtQ), Precipitation of Warmest Quarter (PWmQ), Temperature Annual Range (TAR) and Slope (Slo). These variables are biological significant for the taxon, as they have decisive influence in the critical stages of germination and fruiting. The current and potential distributional areas identified by the model fall mainly in regions with some degree of environmental protection, with some exceptions. A recovery plan for the species should be considered. Species Distribution Modelling cannot substitute long-term monitoring programmes, yet it is a useful tool for identifying appropriate areas of taxon occurrence, and thus allow for efficient use of the economic and human resources.     

The distribution of cultivated species of Porophyllum (Asteraceae) and their wild relatives under climate change

The geographic distribution of plant species is already being affected by climate change. Cropping patterns of edible plant species and their wild relatives will also be affected, making it important to predict possible changes to their distributions in the future. Currently, species distribution models are valuable tools that allow the estimation of species’ potential distributions, in the recent past as well as during other time spans for which climate data have been obtained. With the aim of evaluating how species distributions respond to current and future climate changes, in this work species distribution models were generated for two cultivated species of the Porophyllum genus (Asteraceae), known commonly as ‘pápalos' or ‘pápaloquelites', as well as their Mexican wild relatives, at five points in time (21,000 years ago, present, 2020, 2050, and 2080). Using a database of 1442 entries for 16 species of Porophyllum and 19 environmental variables, species distribution models were constructed for each time period using the Maxent modelling algorithm; those constructed for the future used a severe climate change scenario. The results demonstrate contrasting effects between the two cultivated species; for P. linaria, the future scenario suggests a decrease in distribution area, while for P. macrocephalum distribution is predicted to increase. Similar trends are observed in their wild relatives, where 11 species will tend to decrease in distribution area, while three are predicted to increase. It is concluded that the most important agricultural areas where the cultivated species are grown will not be greatly affected, while the areas inhabited by the wild species will. However, while the results suggest that climate change will affect the distribution of the cultivated species in contrasting ways, evaluations at finer scales are recommended to clarify the impact within cultivation zones.     

Bioclimatic constraints to Andean cat distribution: a modelling application for rare species

Aim To identify the bioclimatic niche of the endangered Andean cat (Leopardus jacobita), one of the rarest and least known felids in the world, by developing a species distribution model. Location South America, High Andes and Patagonian steppe. Peru, Bolivia, Chile, Argentina. Methods We used 108 Andean cat records to build the models, and 27 to test them, applying the Maxent algorithm to sets of uncorrelated bioclimatic variables from global databases, including elevation. We based our biogeographical interpretations on the examination of the predicted geographic range, the modelled response curves and latitudinal variations in climatic variables associated with the locality data. Results Simple bioclimatic models for Andean cats were highly predictive with only 3–4 explanatory variables. The climatic niche of the species was defined by extreme diurnal variations in temperature, cold minimum and moderate maximum temperatures, and aridity, characteristic not only of the Andean highlands but also of the Patagonian steppe. Argentina had the highest representation of suitable climates, and Chile the lowest. The most favourable conditions were centrally located and spanned across international boundaries. Discontinuities in suitable climatic conditions coincided with three biogeographical barriers associated with climatic or topographic transitions. Main conclusions Simple bioclimatic models can produce useful predictions of suitable climatic conditions for rare species, including major biogeographical constraints. In our study case, these constraints are also known to affect the distribution of other Andean species and the genetic structure of Andean cat populations. We recommend surveys of areas with suitable climates and no Andean cat records, including the corridor connecting two core populations. The inclusion of landscape variables at finer scales, crucially the distribution of Andean cat prey, would contribute to refine our predictions for conservation applications.     

Transferable species distribution modelling: Comparative performance of Generalised Functional Response models

Predictive species distribution models (SDMs) are becoming increasingly important in ecology, in the light of rapid environmental change. However, the predictions of most current SDMs are specific to the habitat composition of the environments in which they were fitted. This may limit SDM predictive power because species may respond differently to a given habitat depending on the availability of all habitats in their environment, a phenomenon known as a functional response in resource selection. The Generalised Functional Response (GFR) framework captures this dependence by formulating the SDM coefficients as functions of habitat availability. The original GFR implementation used global polynomial functions of habitat availability to describe the functional responses. In this study, we develop several refinements of this approach and compare their predictive performance using two simulated and two real datasets. We first use local radial basis functions (RBF), a more flexible approach than global polynomials, to represent the habitat selection coefficients, and balance bias with precision via regularization to prevent overfitting. Second, we use the RBF-GFR and GFR models in combination with the classification and regression tree CART, which has more flexibility and better predictive powers for non-linear modelling. As further extensions, we use random forests (RFs) and extreme gradient boosting (XGBoost), ensemble approaches that consistently lead to variance reduction in generalization error. We find that the different methods are ranked consistently across the datasets for out-of-data prediction. The traditional stationary approach to SDMs and the GFR model consistently perform at the bottom of the ranking (simple SDMs underfit, and polynomial GFRs overfit the data). The best methods in our list provide non-negligible improvements in predictive performance, in some cases taking the out-of-sample R2 from 0.3 up to 0.7 across datasets. At times of rapid environmental change and spatial non-stationarity ignoring the effects of functional responses on SDMs, results in two different types of prediction bias (under-prediction or mis-positioning of distribution hotspots). However, not all functional response models perform equally well. The more volatile polynomial GFR models can generate biases through over-prediction. Our results indicate that there are consistently robust GFR approaches that achieve impressive gains in transferability across very different datasets.     

Incorporating spatial autocorrelation into species distribution models alters forecasts of climate‐mediated range shifts

Species distribution models (SDMs) are widely used to forecast changes in the spatial distributions of species and communities in response to climate change. However, spatial autocorrelation (SA) is rarely accounted for in these models, despite its ubiquity in broad‐scale ecological data. While spatial autocorrelation in model residuals is known to result in biased parameter estimates and the inflation of type I errors, the influence of unmodeled SA on species' range forecasts is poorly understood. Here we quantify how accounting for SA in SDMs influences the magnitude of range shift forecasts produced by SDMs for multiple climate change scenarios. SDMs were fitted to simulated data with a known autocorrelation structure, and to field observations of three mangrove communities from northern Australia displaying strong spatial autocorrelation. Three modeling approaches were implemented: environment‐only models (most frequently applied in species' range forecasts), and two approaches that incorporate SA; autologistic models and residuals autocovariate (RAC) models. Differences in forecasts among modeling approaches and climate scenarios were quantified. While all model predictions at the current time closely matched that of the actual current distribution of the mangrove communities, under the climate change scenarios environment‐only models forecast substantially greater range shifts than models incorporating SA. Furthermore, the magnitude of these differences intensified with increasing increments of climate change across the scenarios. When models do not account for SA, forecasts of species' range shifts indicate more extreme impacts of climate change, compared to models that explicitly account for SA. Therefore, where biological or population processes induce substantial autocorrelation in the distribution of organisms, and this is not modeled, model predictions will be inaccurate. These results have global importance for conservation efforts as inaccurate forecasts lead to ineffective prioritization of conservation activities and potentially to avoidable species extinctions.     

Autumn larval cold tolerance does not predict the northern range limit of a widespread butterfly species

Climate change is driving range shifts, and a lack of cold tolerance is hypothesized to constrain insect range expansion at poleward latitudes. However, few, if any, studies have tested this hypothesis during autumn when organisms are subjected to sporadic low‐temperature exposure but may not have become cold‐tolerant yet. In this study, we integrated organismal thermal tolerance measures into species distribution models for larvae of the Giant Swallowtail butterfly, Papilio cresphontes (Lepidoptera: Papilionidae), living at the northern edge of its actively expanding range. Cold hardiness of field‐collected larvae was determined using three common metrics of cold‐induced physiological thresholds: the supercooling point, critical thermal minimum, and survival following cold exposure. P. cresphontes larvae were determined to be tolerant of chilling but generally die at temperatures below their SCP, suggesting they are chill‐tolerant or modestly freeze‐avoidant. Using this information, we examined the importance of low temperatures at a broad scale, by comparing species distribution models of P. cresphontes based only on environmental data derived from other sources to models that also included the cold tolerance parameters generated experimentally. Our modeling revealed that growing degree‐days and precipitation best predicted the distribution of P. cresphontes, while the cold tolerance variables did not explain much variation in habitat suitability. As such, the modeling results were consistent with our experimental results: Low temperatures in autumn are unlikely to limit the distribution of P. cresphontes. Understanding the factors that limit species distributions is key to predicting how climate change will drive species range shifts.     

Ice age distributions of European small mammals: insights from species distribution modelling

Aim In addition to the traditionally recognized Last Glacial Maximum (LGM, 21 ka) refuge areas in the Mediterranean region, more northerly LGM distributions for temperate and boreal taxa in central and eastern Europe are increasingly being discussed based on palaeoecological and phylogeographical evidence. Our aim was to investigate the potential refuge locations using species distribution modelling to estimate the geographical distribution of suitable climatic conditions for selected rodent species during the LGM. Location Eurasia. Methods Presence/absence data for seven rodent species with range limits corresponding to the limits of temperate or boreal forest or arctic tundra were used in the analysis. We developed predictive distribution models based on the species present‐day European distributions and validated these against their present‐day Siberian ranges. The models with the best predictors of the species distributions across Siberia were projected onto LGM climate simulations to assess the distribution of climatically suitable areas. Results The best distribution models provided good predictions of the present‐day Siberian ranges of the study species. Their LGM projections showed that areas with a suitable LGM climate for the three temperate species (Apodemus flavicollis, Apodemus sylvaticus and Microtus arvalis) were largely restricted to the traditionally recognized southern refuge areas, i.e. mainly in the Mediterranean region, but also southernmost France and southern parts of the Russian Plain. In contrast, suitable climatic conditions for the two boreal species (Clethrionomys glareous and Microtus agrestis) were predicted as far north as southern England and across southern parts of central and eastern Europe eastwards into the Russian Plain. For the two arctic species (Lemmus lemmus and Microtus oeconomus), suitable climate was predicted from the Atlantic coast eastward across central Europe and into Russia. Main conclusions Our results support the idea of more northerly refuge areas in Europe, indicating that boreal species would have found suitable living conditions over much of southern central and eastern Europe and the Russian Plain. Temperate species would have primarily found suitable conditions in the traditional southern refuge areas, but interestingly also in much of the southern Russian Plain.     

Delimiting the geographical background in species distribution modelling

Aim The extent of the study area (geographical background, GB) can strongly affect the results of species distribution models (SDMs), but as yet we lack objective and practicable criteria for delimiting the appropriate GB. We propose an approach to this problem using trend surface analysis (TSA) and provide an assessment of the effects of varying GB extent on the performance of SDMs for four species. Location Mainland Spain. Methods Using data for four well known wild ungulate species and different GBs delimited with a TSA, we assessed the effects of GB extent on the predictive performance of SDMs: specifically on model calibration (Miller’s statistic) and discrimination (area under the curve of the receiver operating characteristic plot, AUC; sensitivity and specificity), and on the tendency of the models to predict environmental potential when they are projected beyond their training area. Results In the training area, discrimination significantly increased and calibration decreased as the GB was enlarged. In contrast, as GB was enlarged, both discriminatory power and calibration decreased when assessed in the core area of the species distributions. When models trained using small GBs were projected beyond their training area, they showed a tendency to predict higher environmental potential for the species than those models trained using large GBs. Main conclusions By restricting GB extent using a geographical criterion, model performance in the core area of the species distribution can be significantly improved. Large GBs make models demonstrate high discriminatory power but are barely informative. By delimiting GB using a geographical criterion, the effect of historical events on model parameterization may be reduced. Thus purely environmental models are obtained that, when projected onto a new scenario, depict the potential distribution of the species. We therefore recommend the use of TSA in geographically delimiting the GB for use in SDMs.