排序和广义线性模型与广义可加模型在植物种与环境关系研究中的应用

排序和广义线性模型(Generalized Linear Model,GLM)与广义可加模型(Goneralized Additive Model,GAM)是研究植物种与环境间关系的重要方法。基于线性模型的排序方法应限定于环境梯度较短的植被数据。而基于单峰模型的排序方法更适用于梯度较长的情况。PCA、CA／RA系列和CCA系列是常用的排序方法。同时进行环境数据和植被数据分析的CCA系列,能清楚地得出植物种与环境间的关系。CCA改进后的DCCA和PCCA,是现今较理想的排序方法。GLM和GAM实质上是用环境变量的高阶多项式来拟合植物种与环境变量的关系。GLM和GAM扩展了植物种与环境变量之间的关系模型,能深入地探讨植物种与环境间的关系。GLM主要是模型决定的,而GAM主要取决于原始数据。一般来说,排序能得出研究区域的主要环境梯度,提供了物种聚集和植物群落的概略描述。GLM与GAM对于深入研究单个植物种与环境间的关系具有优势。在实际研究中,两种方法结合使用能互补不足。     

Evaluating the ecological realism of plant species distribution models with ecological indicator values

Species distribution models (SDMs) are routinely applied to assess current as well as future species distributions, for example to assess impacts of future environmental change on biodiversity or to underpin conservation planning. It has been repeatedly emphasized that SDMs should be evaluated based not only on their goodness of fit to the data, but also on the realism of the modeled ecological responses. However, possibilities for the latter are hampered by limited knowledge on the true responses as well as a lack of quantitative evaluation methods. Here we compared modeled niche optima obtained from European-scale SDMs of 1476 terrestrial vascular plant species with empirical ecological indicator values indicating the preferences of plant species for key environmental conditions. For each plant species we first fitted an ensemble SDM including three modeling techniques (GLM, GAM and BRT) and extracted niche optima for climate, soil, land use and nitrogen deposition variables with a large explanatory power for the occurrence of that species. We then compared these SDM-derived niche optima with the ecological indicator values by means of bivariate correlation analysis. We found weak to moderate correlations in the expected direction between the SDM-derived niche optima and ecological indicator values. The strongest correlation occurred between the modeled optima for growing degree days and the ecological indicator values for temperature. Correlations were weaker for SDM-derived niche optima with a more distal relationship to ecological indicator values (notably precipitation and soil moisture). Further, correlations were consistently highest for BRT, followed by GLM and GAM. Our method gives insight into the ecological realism of modeled niche optima and projected core habitats and can be used to improve SDMs by making a more informed selection of environmental variables and modeling techniques.     

Land use improves spatial predictions of mountain plant abundance but not presence‐absence

Question: Does a land‐use variable improve spatial predictions of plant species presence‐absence and abundance models at the regional scale in a mountain landscape? Location: Western Swiss Alps. Methods: Presence‐absence generalized linear models (GLM) and abundance ordinal logistic regression models (LRM) were fitted to data on 78 mountain plant species, with topo‐climatic and/or land‐use variables available at a 25‐m resolution. The additional contribution of land use when added to topo‐climatic models was evaluated by: (1) assessing the changes in model fit and (2) predictive power, (3) partitioning the deviance respectively explained by the topo‐climatic variables and the land‐use variable through variation partitioning, and (5) comparing spatial projections. Results: Land use significantly improved the fit of presence‐absence models but not their predictive power. In contrast, land use significantly improved both the fit and predictive power of abundance models. Variation partitioning also showed that the individual contribution of land use to the deviance explained by presence‐absence models was, on average, weak for both GLM and LRM (3.7% and 4.5%, respectively), but changes in spatial projections could nevertheless be important for some species. Conclusions: In this mountain area and at our regional scale, land use is important for predicting abundance, but not presence‐absence. The importance of adding land‐use information depends on the species considered. Even without a marked effect on model fit and predictive performance, adding land use can affect spatial projections of both presence‐absence and abundance models.     

A comparison of absolute performance of different correlative and mechanistic species distribution models in an independent area

To investigate the comparative abilities of six different bioclimatic models in an independent area, utilizing the distribution of eight different species available at a global scale and in Australia. Global scale and Australia. We tested a variety of bioclimatic models for eight different plant species employing five discriminatory correlative species distribution models (SDMs) including Generalized Linear Model (GLM), MaxEnt, Random Forest (RF), Boosted Regression Tree (BRT), Bioclim, together with CLIMEX (CL) as a mechanistic niche model. These models were fitted using a training dataset of available global data, but with the exclusion of Australian locations. The capabilities of these techniques in projecting suitable climate, based on independent records for these species in Australia, were compared. Thus, Australia is not used to calibrate the models and therefore it is as an independent area regarding geographic locations. To assess and compare performance, we utilized the area under the receiver operating characteristic (ROC) curves (AUC), true skill statistic (TSS), and fractional predicted areas for all SDMs. In addition, we assessed satisfactory agreements between the outputs of the six different bioclimatic models, for all eight species in Australia. The modeling method impacted on potential distribution predictions under current climate. However, the utilization of sensitivity and the fractional predicted areas showed that GLM, MaxEnt, Bioclim, and CL had the highest sensitivity for Australian climate conditions. Bioclim calculated the highest fractional predicted area of an independent area, while RF and BRT were poor. For many applications, it is difficult to decide which bioclimatic model to use. This research shows that variable results are obtained using different SDMs in an independent area. This research also shows that the SDMs produce different results for different species; for example, Bioclim may not be good for one species but works better for other species. Also, when projecting a “large” number of species into novel environments or in an independent area, the selection of the “best” model/technique is often less reliable than an ensemble modeling approach. In addition, it is vital to understand the accuracy of SDMs' predictions. Further, while TSS, together with fractional predicted areas, are appropriate tools for the measurement of accuracy between model results, particularly when undertaking projections on an independent area, AUC has been proved not to be. Our study highlights that each one of these models (CL, Bioclim, GLM, MaxEnt, BRT, and RF) provides slightly different results on projections and that it may be safer to use an ensemble of models.     

Are niche‐based species distribution models transferable in space?

Aim To assess the geographical transferability of niche‐based species distribution models fitted with two modelling techniques. Location Two distinct geographical study areas in Switzerland and Austria, in the subalpine and alpine belts. Methods Generalized linear and generalized additive models (GLM and GAM) with a binomial probability distribution and a logit link were fitted for 54 plant species, based on topoclimatic predictor variables. These models were then evaluated quantitatively and used for spatially explicit predictions within (internal evaluation and prediction) and between (external evaluation and prediction) the two regions. Comparisons of evaluations and spatial predictions between regions and models were conducted in order to test if species and methods meet the criteria of full transferability. By full transferability, we mean that: (1) the internal evaluation of models fitted in region A and B must be similar; (2) a model fitted in region A must at least retain a comparable external evaluation when projected into region B, and vice‐versa; and (3) internal and external spatial predictions have to match within both regions. Results The measures of model fit are, on average, 24% higher for GAMs than for GLMs in both regions. However, the differences between internal and external evaluations (AUC coefficient) are also higher for GAMs than for GLMs (a difference of 30% for models fitted in Switzerland and 54% for models fitted in Austria). Transferability, as measured with the AUC evaluation, fails for 68% of the species in Switzerland and 55% in Austria for GLMs (respectively for 67% and 53% of the species for GAMs). For both GAMs and GLMs, the agreement between internal and external predictions is rather weak on average (Kulczynski's coefficient in the range 0.3–0.4), but varies widely among individual species. The dominant pattern is an asymmetrical transferability between the two study regions (a mean decrease of 20% for the AUC coefficient when the models are transferred from Switzerland and 13% when they are transferred from Austria). Main conclusions The large inter‐specific variability observed among the 54 study species underlines the need to consider more than a few species to test properly the transferability of species distribution models. The pronounced asymmetry in transferability between the two study regions may be due to peculiarities of these regions, such as differences in the ranges of environmental predictors or the varied impact of land‐use history, or to species‐specific reasons like differential phenotypic plasticity, existence of ecotypes or varied dependence on biotic interactions that are not properly incorporated into niche‐based models. The lower variation between internal and external evaluation of GLMs compared to GAMs further suggests that overfitting may reduce transferability. Overall, a limited geographical transferability calls for caution when projecting niche‐based models for assessing the fate of species in future environments.     

BIOMOD – optimizing predictions of species distributions and projecting potential future shifts under global change

A new computation framework (BIOMOD: BIOdiversity MODelling) is presented, which aims to maximize the predictive accuracy of current species distributions and the reliability of future potential distributions using different types of statistical modelling methods. BIOMOD capitalizes on the different techniques used in static modelling to provide spatial predictions. It computes, for each species and in the same package, the four most widely used modelling techniques in species predictions, namely Generalized Linear Models (GLM), Generalized Additive Models (GAM), Classification and Regression Tree analysis (CART) and Artificial Neural Networks (ANN). BIOMOD was applied to 61 species of trees in Europe using climatic quantities as explanatory variables of current distributions. On average, all the different modelling methods yielded very good agreement between observed and predicted distributions. However, the relative performance of different techniques was idiosyncratic across species, suggesting that the most accurate model varies between species. The results of this evaluation also highlight that slight differences between current predictions from different modelling techniques are exacerbated in future projections. Therefore, it is difficult to assess the reliability of alternative projections without validation techniques or expert opinion. It is concluded that rather than using a single modelling technique to predict the distribution of several species, it would be more reliable to use a framework assessing different models for each species and selecting the most accurate one using both evaluation methods and expert knowledge.     

Environmental control of plant species abundance in a microtidal Mediterranean saltmarsh

Question: To what extent are environmental factors the main determinants of species abundance in Mediterranean coastal marshlands? Location: The Llobregat delta, Barcelona, Spain. Methods: Vegetation relevés were performed and a set of water table and soil variables were periodically monitored in 43 sampling points randomly distributed in four marsh areas (sites) along a coastal–inland gradient. A canonical correspondence analysis (CCA) was performed to identify the primary water and soil correlates of species cover, after considering the effect of site and point spatial location. The realized niches of dominant species were modeled through GLMs performed on the first two axes of CCA. Niche overlapping among these species was compared with their coexistence, assessed through pairwise correlations of relative species cover in each sampling point. Results: Water and soil variables explained more of the variation in species' abundance than site and spatial position. Mean water table level, maximum water conductivity and sodium adsorption ratio (SAR), summarized in the two first CCA axes, explained 23.8% of the variability in species' cover. Arthrocnemum fruticosum, Phragmites australis subsp. australis, Juncus acutus, Spartina versicolor and Juncus maritimus dominated the vegetation stands. Niches obtained from GLM response curves showed moderate overlapping among all these species except for A. fruticosum. However, pairwise correlations were mainly negative or non‐significant, indicating low coincidence, and even segregation, between species' cover. Conclusions: The abundance of dominant plants in Mediterranean marshes is only partly explained by the environmental gradients summarized in niche models. The role of other factors such as facilitation or competition between species and random recruitment should be explored.     

Geomorphological disturbance is necessary for predicting fine‐scale species distributions

Disturbances related to geomorphological processes are frequent, widespread and often intense at high latitudes and altitudes, affecting the fine‐scale distribution of many plant species. While the inclusion of physical disturbances into models of species geographic ranges is widely recommended, no studies have yet tested the utility of field‐quantified geomorphological disturbances for terrestrial species distribution modelling. Here we apply generalized additive models and boosted regression trees to examine if the explicit inclusion of terrestrial and fluvial geomorphological variables alters species distribution models for 154 vascular plant, bryophyte and lichen species in north European mountain tundra. The inclusion of these disturbances significantly improved both the explanatory and predictive power of distribution models, with consistent results for all three species groups. Spatial distribution predictions changed considerably for some species after the inclusion of disturbance variables, with fluvial disturbances generating strongly linear features for species influenced by erosion or sediment deposition. As a consequence, models incorporating geomorphological variables produced markedly more refined distribution maps than simpler models. Predictions of species distributions will thus benefit strongly from the inclusion of fine‐scale geomorphological variables, particularly in areas of active earth surface processes, enabling more accurate forecasting of future species ranges under changing conditions.     

Enhanced effects of biotic interactions on predicting multispecies spatial distribution of submerged macrophytes after eutrophication

Water eutrophication creates unfavorable environmental conditions for submerged macrophytes. In these situations, biotic interactions may be particularly important for explaining and predicting the submerged macrophytes occurrence. Here, we evaluate the roles of biotic interactions in predicting spatial occurrence of submerged macrophytes in 1959 and 2009 for Dianshan Lake in eastern China, which became eutrophic since the 1980s. For the four common species occurred in 1959 and 2009, null species distribution models based on abiotic variables and full models based on both abiotic and biotic variables were developed using generalized linear model (GLM) and boosted regression trees (BRT) to determine whether the biotic variables improved the model performance. Hierarchical Bayesian‐based joint species distribution models capable of detecting paired biotic interactions were established for each species in both periods to evaluate the changes in the biotic interactions. In most of the GLM and BRT models, the full models showed better performance than the null models in predicting the species presence/absence, and the relative importance of the biotic variables in the full models increased from less than 50% in 1959 to more than 50% in 2009 for each species. Moreover, co‐occurrence correlation of each paired species interaction was higher in 2009 than that in 1959. The findings suggest biotic interactions that tend to be positive play more important roles in the spatial distribution of multispecies assemblages of macrophytes and should be included in prediction models to improve prediction accuracy when forecasting macrophytes’ distribution under eutrophication stress.     

Biotic predictors with phenological information improve range estimates for migrating monarch butterflies in Mexico

Although long-standing theory suggests that biotic variables are only relevant at local scales for explaining the patterns of species' distributions, recent studies have demonstrated improvements to species distribution models (SDMs) by incorporating predictor variables informed by biotic interactions. However, some key methodological questions remain, such as which kinds of interactions are permitted to include in these models, how to incorporate the effects of multiple interacting species, and how to account for interactions that may have a temporal dependence. We addressed these questions in an effort to model the distribution of the monarch butterfly Danaus plexippus during its fall migration (September–November) through Mexico, a region with new monitoring data and uncertain range limits even for this well-studied insect. We estimated species richness of selected nectar plants (Asclepias spp.) and roosting trees (various highland species) for use as biotic variables in our models. To account for flowering phenology, we additionally estimated nectar plant richness of flowering species per month. We evaluated three types of models: climatic variables only (abiotic), plant richness estimates only (biotic) and combined (abiotic and biotic). We selected models with AICc and additionally determined if they performed better than random on spatially withheld data. We found that the combined models accounting for phenology performed best for all three months, and better than random for discriminatory ability but not omission rate. These combined models also produced the most ecologically realistic spatial patterns, but the modeled response for nectar plant richness matched ecological predictions for November only. These results represent the first model-based monarch distributional estimates for the Mexican migration route and should provide foundations for future conservation work. More generally, the study demonstrates the potential benefits of using SDM-derived richness estimates and phenological information for biotic factors affecting species distributions.     

Bagging GLM: Improved generalized linear model for the analysis of zero-inflated data

Species-occurrence data sets tend to contain a large proportion of zero values, i.e., absence values (zero-inflated). Statistical inference using such data sets is likely to be inefficient or lead to incorrect conclusions unless the data are treated carefully. In this study, we propose a new modeling method to overcome the problems caused by zero-inflated data sets that involves a regression model and a machine-learning technique. We combined a generalized liner model (GLM), which is widely used in ecology, and bootstrap aggregation (bagging), a machine-learning technique. We established distribution models of Vincetoxicum pycnostelma (a vascular plant) and Ninox scutulata (an owl), both of which are endangered and have zero-inflated distribution patterns, using our new method and traditional GLM and compared model performances. At the same time we modeled four theoretical data sets that contained different ratios of presence/absence values using new and traditional methods and also compared model performances. For distribution models, our new method showed good performance compared to traditional GLMs. After bagging, area under the curve (AUC) values were almost the same as with traditional methods, but sensitivity values were higher. Additionally, our new method showed high sensitivity values compared to the traditional GLM when modeling a theoretical data set containing a large proportion of zero values. These results indicate that our new method has high predictive ability with presence data when analyzing zero-inflated data sets. Generally, predicting presence data is more difficult than predicting absence data. Our new modeling method has potential for advancing species distribution modeling.     

Predicting the species composition of Nardus stricta communities by logistic regression modelling

Question: Predictive models in plant ecology usually deal with single species or community types. Little effort has so far been made to predict the species composition of a community explicitly. The modelling approach presented here provides a conceptual framework on how to achieve this by combining habitat models for a large number of species to an additive community model. Our approach is exemplified by Nardus stricta communities (acidophilous, low‐productive grassland). Location: Large areas of Germany, 0–2040 m a.s.l. Methods: Logistic regression is applied for individual species models which are subsequently combined for an explicit prediction of species composition. Several parameters reflecting soil, management and climatic conditions serve as predictor variables. For validation, bootstrap and jackknife resampling procedures are used as well as ordination techniques (DCA, CCA). Results: We calculated significant models for 138 individual species. The predictions of species composition and species richness yield good agreements with the observed data. DCA and CCA results show that the community model preserves the main patterns in floristic space. Conclusions: Our approach of predicting species composition is an effective tool that can be applied in nature conservation, e.g. to assess the effects of different site conditions and alternative management scenarios on species composition and richness.     

Predicting species distributions: a critical comparison of the most common statistical models using artificial species

Aim To test statistical models used to predict species distributions under different shapes of occurrence–environment relationship. We addressed three questions: (1) Is there a statistical technique that has a consistently higher predictive ability than others for all kinds of relationships? (2) How does species prevalence influence the relative performance of models? (3) When an automated stepwise selection procedure is used, does it improve predictive modelling, and are the relevant variables being selected? Location We used environmental data from a real landscape, the state of California, and simulated species distributions within this landscape. Methods Eighteen artificial species were generated, which varied in their occurrence response to the environmental gradients considered (random, linear, Gaussian, threshold or mixed), in the interaction of those factors (no interaction vs. multiplicative), and on their prevalence (50% vs. 5%). The landscape was then randomly sampled with a large (n = 2000) or small (n = 150) sample size, and the predictive ability of each statistical approach was assessed by comparing the true and predicted distributions using five different indexes of performance (area under the receiver‐operator characteristic curve, Kappa, correlation between true and predictive probability of occurrence, sensitivity and specificity). We compared generalized additive models (GAM) with and without flexible degrees of freedom, logistic regressions (general linear models, GLM) with and without variable selection, classification trees, and the genetic algorithm for rule‐set production (GARP). Results Species with threshold and mixed responses, additive environmental effects, and high prevalence generated better predictions than did other species for all statistical models. In general, GAM outperforms all other strategies, although differences with GLM are usually not significant. The two variable‐selection strategies presented here did not discriminate successfully between truly causal factors and correlated environmental variables. Main conclusions Based on our analyses, we recommend the use of GAM or GLM over classification trees or GARP, and the specification of any suspected interaction terms between predictors. An expert‐based variable selection procedure was preferable to the automated procedures used here. Finally, for low‐prevalence species, variability in model performance is both very high and sample‐dependent. This suggests that distribution models for species with low prevalence can be improved through targeted sampling.     

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 forecasts.     

Generalized models vs. classification tree analysis: Predicting spatial distributions of plant species at different scales

Abstract. Statistical models of the realized niche of species are increasingly used, but systematic comparisons of alternative methods are still limited. In particular, only few studies have explored the effect of scale in model outputs. In this paper, we investigate the predictive ability of three statistical methods (generalized linear models, generalized additive models and classification tree analysis) using species distribution data at three scales: fine (Catalonia), intermediate (Portugal) and coarse (Europe). Four Mediterranean tree species were modelled for comparison. Variables selected by models were relatively consistent across scales and the predictive accuracy of models varied only slightly. However, there were slight differences in the performance of methods. Classification tree analysis had a lower accuracy than the generalized methods, especially at finer scales. The performance of generalized linear models also increased with scale. At the fine scale GLM with linear terms showed better accuracy than GLM with quadratic and polynomial terms. This is probably because distributions at finer scales represent a linear sub‐sample of entire realized niches of species. In contrast to GLM, the performance of GAM was constant across scales being more data‐oriented. The predictive accuracy of GAM was always at least equal to other techniques, suggesting that this modelling approach is more robust to variations of scale because it can deal with any response shape.     

Do Stacked Species Distribution Models Reflect Altitudinal Diversity Patterns?

The objective of this study was to evaluate the performance of stacked species distribution models in predicting the alpha and gamma species diversity patterns of two important plant clades along elevation in the Andes. We modelled the distribution of the species in the Anthurium genus (53 species) and the Bromeliaceae family (89 species) using six modelling techniques. We combined all of the predictions for the same species in ensemble models based on two different criteria: the average of the rescaled predictions by all techniques and the average of the best techniques. The rescaled predictions were then reclassified into binary predictions (presence/absence). By stacking either the original predictions or binary predictions for both ensemble procedures, we obtained four different species richness models per taxa. The gamma and alpha diversity per elevation band (500 m) was also computed. To evaluate the prediction abilities for the four predictions of species richness and gamma diversity, the models were compared with the real data along an elevation gradient that was independently compiled by specialists. Finally, we also tested whether our richness models performed better than a null model of altitudinal changes of diversity based on the literature. Stacking of the ensemble prediction of the individual species models generated richness models that proved to be well correlated with the observed alpha diversity richness patterns along elevation and with the gamma diversity derived from the literature. Overall, these models tend to overpredict species richness. The use of the ensemble predictions from the species models built with different techniques seems very promising for modelling of species assemblages. Stacking of the binary models reduced the over-prediction, although more research is needed. The randomisation test proved to be a promising method for testing the performance of the stacked models, but other implementations may still be developed.     

Integrating Multiple Distribution Models to Guide Conservation Efforts of an Endangered Toad

Species distribution models are used for numerous purposes such as predicting changes in species’ ranges and identifying biodiversity hotspots. Although implications of distribution models for conservation are often implicit, few studies use these tools explicitly to inform conservation efforts. Herein, we illustrate how multiple distribution models developed using distinct sets of environmental variables can be integrated to aid in identification sites for use in conservation. We focus on the endangered arroyo toad (Anaxyrus californicus), which relies on open, sandy streams and surrounding floodplains in southern California, USA, and northern Baja California, Mexico. Declines of the species are largely attributed to habitat degradation associated with vegetation encroachment, invasive predators, and altered hydrologic regimes. We had three main goals: 1) develop a model of potential habitat for arroyo toads, based on long-term environmental variables and all available locality data; 2) develop a model of the species’ current habitat by incorporating recent remotely-sensed variables and only using recent locality data; and 3) integrate results of both models to identify sites that may be employed in conservation efforts. We used a machine learning technique, Random Forests, to develop the models, focused on riparian zones in southern California. We identified 14.37% and 10.50% of our study area as potential and current habitat for the arroyo toad, respectively. Generally, inclusion of remotely-sensed variables reduced modeled suitability of sites, thus many areas modeled as potential habitat were not modeled as current habitat. We propose such sites could be made suitable for arroyo toads through active management, increasing current habitat by up to 67.02%. Our general approach can be employed to guide conservation efforts of virtually any species with sufficient data necessary to develop appropriate distribution models.     

Modelling and predicting fungal distribution patterns using herbarium data

Aim The main aims of this study are: (1) to test if temperature and related parameters are the primary determinants of the regional distribution of macrofungi (as is commonly recognized for plants); (2) to test if the success of modelling fungal distribution patterns depends on species and distribution characteristics; and (3) to explore the potential of using herbarium data for modelling and predicting fungal species’ distributions. Location The study area, Norway, spans 58–71° N latitude and 4–32° E longitude, and embraces extensive ecological gradients in a small area. Methods The study is based on 1020 herbarium collections of nine selected species of macrofungi and a set of 75 environmental predictor variables, all recorded in a 5 × 5‐km grid covering Norway. Primarily, generalized linear model (GLM; logistic regression) analyses were used to identify the environmental variables that best accounted for the species’ recorded distributions in Norway. Second, Maxent analyses (using variables identified by GLM) were used to produce predictive potential distribution maps for these species. Results Variables relating to temperature and radiation were most frequently included in the GLMs, and between 24.8% and 59.8% of the variation in single‐species occurrence was accounted for. The fraction of variation explained by the GLMs ranged from 41.6% to 59.8% for species with restricted distributions, and from 24.8% to 39.3% for species with widespread/scattered and intermediate distributions. The two‐step procedure of GLM followed by Maxent gave predictions with very high values for the area under the curve (0.927–0.997), and maps of potential distribution were generally credible. Main conclusions We show that temperature is a key factor governing the distribution of macrofungi in Norway, indicating that fungi may respond strongly to global warming. We confirm that modelling success depends partly on species and distribution characteristics, notably on how the distribution relates to the extent of the study area. Our study demonstrates that the combination of GLM and Maxent may be a fruitful approach for biogeography. We conclude that herbarium data improve insight into factors that control the distributions of fungi, of particular value for research on fleshy fungi (mushrooms), which have largely cryptic life cycles.     

气候变化对物种分布影响模拟中的不确定性组分分割与制图——以油松为例

物种分布模型是预测评估气候变化对物种分布影响的主要工具。为了降低物种分布模型在预测过程中的不确定性,近期有学者提出了采用组合预测的新方法,即采用多套建模数据、模型技术,模型参数,以及环境情景数据对物种分布进行预测,构成物种分布预测集合。但是,组合预测中各组分对变异的贡献还知之甚少,因此有必要把变异组分来源进行分割,以更有效地利用组合预测方法来降低模型预测中的不确定性。以油松为例,采用8个生态位模型,9套模型训练数据,3个GCM模型和一个SRES(A2)排放情景,模型分析了油松当前(1961-1990年)和未来气候条件下3个时间段(2010-2039年,2040-2069年,2070-2099年)的潜在分布。共计得到当前分布预测数据72套,未来每个时间段分布数据216套。采用开发的ClimateChina软件进行当前和未来气候数据的降尺度处理。采用Kappa、真实技巧统计方法(TSS)和接收机工作特征曲线下的面积(AUC)对模型预测能力进行评估。结果表明,随机森林(RF)、广义线性模型(GLM),广义加法模型(GAM)、多元自适应样条函数(MARS)以及助推法(GBM)预测效果较好,几乎不受建模数据之间差异的影响。混合判别分析模型(MDA)对建模数据之间的差异非常敏感,甚至出现建模失败现象。采用三因素方差分析方法对组合预测中的不确定性来源进行变异分割,结果表明,模型之间的差异对模拟预测结果不确定性的贡献最大且所占比例极高,而建模数据之间的差异贡献最小,GCM贡献居中。研究将有助于加深对物种分布模拟预测中不确定性的认识。