物种分布模型的发展及评价方法

物种分布模型已被广泛地应用于以保护区规划、气候变化对物种分布的影响等为目的的研究。回顾了已经得到广泛应用的多种物种分布模型,总结了评价模型性能的方法。基于物种分布模型的发展和应用以及性能评价中尚存在的问题,本文认为:在物种分布模型中集成样本选择模块能够避免模型预测过程中的过度拟合及欠拟合,增加变量选择模块可评估和降低变量之间自相关性的影响,增加生物因子以及将物种对环境的适应性机制(及扩散行为特征)和潜在分布模型进行结合,是提高模型预测性能的可行方案;在模型性能的评价方面,采用赤池信息量可对模型的预测性能进行客观评价。相关建议可为物种分布建模提供参考。     

基于集成模型的小黄鱼越冬群体适宜生境及其环境影响因素

小黄鱼是中韩渔业共同利用鱼种,其跨界洄游习性限制了对越冬场范围的调查和评估,导致对越冬群体适宜栖息地分布缺乏了解。本研究基于越冬期我国自然海域的物种分布点位数据和5个环境数据,运用8个物种分布模型(SDM)分析了小黄鱼越冬场分布范围,采用5折交叉验证,利用受试者工作特征曲线下面积(AUC)评价模型预测性能,并通过加权集成方法构建综合生境模型预测越冬场核心分布位置。结果表明: 出现/未出现数据模型预测准确度普遍高于仅出现模型;在出现/未出现数据模型中,机器学习方法预测准确度高于经典回归模型,支持向量模型(SVM)准确度最高(AUC=0.85),广义线性模型(GLM)准确度最低(AUC=0.73)。集成模型AUC较单一独立模型的准确度有所提升,表明集成模型能有效降低单一独立模型所带来的不确定性,提高模型预测准确度。变量重要性分析结果显示,盐度和温度是决定小黄鱼越冬场地理分布的重要因素,适宜分布区集中在黄海南部外海、东海北部外海和浙江省沿岸海域,而黄海南部沿岸海域和东海中南部外海为不适宜越冬区。研究结果为预测小黄鱼潜在越冬场提供了理论基础,可支撑越冬场渔业资源的空间规划和可持续利用。     

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。     

气候变化条件下太平洋丽龟的适宜生境及其变化

太平洋丽龟作为被国际自然保护联盟(IUCN)认定的脆弱物种,近年来备受关注。为了解当前及未来气候情景条件下太平洋丽龟的分布及其变化,本研究利用其发现记录和8个环境预测变量(包括深度、离岸距离、平均初级生产力、最小初级生产力、海表平均温度、海表最小温度、海表平均盐度、海表最小盐度),构建了组合物种分布模型(Ensemble SDM)对其潜在栖息地分布进行预测,并利用曲线下面积(AUC)和真实技巧统计(TSS)值评估模型的准确性。结果表明：AUC和TSS值分别为0.96和0.81,表明组合模型具有较好的预测性能;海洋表面温度和盐度是决定太平洋丽龟分布最重要的两个预测变量,适宜温度为23～29℃,适宜盐度<34;当前环境条件下太平洋丽龟分布范围在30°N—25°S;在未来气候情景条件下,该物种的分布范围将减少,特别是在2100s RCP85气候情境下,其适宜生存范围将减少28%。模型验证结果显示,模型准确性较高,能对太平洋丽龟在当前和未来气候情景下的分布做出较为准确的预测。本研究可为制定更加合理的保护措施和管理策略提供数据参考。     

样条变换偏最小二乘在肝癌数据分类中的应用

肝癌是中国最常见的恶性肿瘤之一。基于肿瘤基因表达谱数据的分析与研究是当今研究的热点,对于癌症的早期诊断、治疗具有十分重要的意义。针对高维小样本基因表达谱数据所显现的变量间严重共线性、类别变量与预测变量的非线性关系,采用了基于样条变换的偏最小二乘回归新技术。首先通过筛选法去除基因表达谱数据中的冗余信息,然后以3次B基样条变换实现非线性基因表达谱数据的线性化重构,随后将重构的矩阵交由偏最小二乘法构建类别变量与预测变量间的关系模型。最后,通过对肝癌肿瘤基因表达谱数据的分析,结果显示此分类模型对数据重构稳健,有效的解决了高维小样本基因表达谱数据间的过拟合和变量间的共线性,具有较高的拟合和分类正确率。     

人参潜在地理分布以及气候变化对其影响预测

本文以人参为研究对象,基于人参分布点位数据和22个气候环境因子数据,运用BioMod2平台10个物种分布模型对当前我国东北地区人参潜在生境分布进行预测.以受试者工作特征曲线(ROC)为权重集成10个模型的模拟结果,构建组合模型,并基于该模型预测了IPCC 第五次评估报告中RCP 8.5、RCP 6.0、RCP 4.5和RCP 2.6等4种排放情景下21世纪50和70年代人参潜在分布范围.结果表明: 在基准气候条件下,人参适宜生境面积占研究区总面积的10.4%,此类地区主要分布于研究区东北部长白山地区以及小兴安岭东南部区域的森林地带.在未来不同的排放情景下研究区人参的适宜生境变化显著,总体上分布范围将有一定程度的缩小.同时参与建模的10种模型在统计学精度、预测结果以及变量权重上都有差异.模型精度计算结果表明,MAXENT模拟效果最好,GAM、RF和ANN次之,SRE模拟精度最低.本文构建的组合模型在一定程度上提高了现有物种分布模型的预测精度,从而使模拟效果更优.     

样本量与研究范围变化对MaxEnt模型准确度的影响——以黑白仰鼻猴为例

MaxEnt模型是过去几年最为流行的物种分布预测模型之一。针对一些濒危物种、入侵种和模拟数据的研究表明,MaxEnt模型均能在小样本的分布数据下得到较准确的预测结果。此外,研究范围的变化也会影响MaxEnt模型的构建。 然而,基于动物的实际分布数据来评估MaxEnt模型的研究甚少。 我们以黑白仰鼻猴 (Rhinopithecus bieti)为例,以11个猴群的分布数据为训练数据（样本量从1到10个猴群）,在不同研究范围内构建MaxEnt模型,通过其它5个的猴群分布数据验证,分析样本量和研究范围变化对模型准确度产生的影响。 结果表明,随样本量和研究范围增大,MaxEnt模型准确度及稳定性都有增加。 此外,研究范围变化对模型准确度有一定影响。 应用Maxent进行物种分布预测时,训练数据应尽可能涵盖该物种可能出现的全部环境梯度。构建模型所需的背景数据点选择,应与建模使用的物种出现点形成有效对照。     

美国大陆外来入侵物种斑马纹贻贝(Dreissena polymorpha)潜在生境预测模型

防止外来生物入侵造成危害的重要手段是阻止可能造成入侵的物种进入适合其生存的地区.论文以1864个美国外来入侵物种斑马纹贻贝定点发生数据和开放式基础地理信息数据库Daymet的34个环境变量为主要信息源,采用逻辑斯蒂回归(LR)、分类与回归树模型(CART)、基于规则的遗传算法(GARP)、最大熵法(Maxent)4种途径,建立美国大陆部分潜在生境预测模型,从接受者运行特征曲线下面积(AUC)、Pearson相关系数、Kappa值3个方面来检验模型预测精度,在此基础上分析斑马纹贻贝的空间分布规律及其环境影响因素.研究结果表明:在3个评价指标中,4个生态位模型预测精度均达到优良水平,其中Maxent在物种现实生境模拟、主要生态环境因子筛选、环境因子对物种生境影响的定量描述方面都表现出了优越的性能;距水源距离、海拔高度、降水频率、太阳辐射是影响物种空间分布的主要环境因子.论文提出的研究方法对中国外来入侵物种生境预测具有较强的借鉴意义,研究结果对中国海洋外来入侵物种沙筛贝的预测与防治,具有一定的指导作用.     

PSDS 2.0: 一个基于GIS和多个模型的生物潜在分布地预测系统

根据对生物分布地预测模型和软件发展现状的分析和总结, 本研究在PSDS 1.0的基础上提出并实现一个基于GIS且具有多个代表性模型的生物分布地预测系统(PSDS 2.0)。PSDS 2.0系统继承了1.0的环境包络和聚类包络模型, 进一步引入了限制因子包络、马氏距离、支持向量机等新模型, 并针对本领域中模型比较与选择的难点增加了迭代交叉验证的多模型选择功能。系统还实现了灵活定制和评估伪负样本的功能, 通过用只需要正样本的I类模型预测的结果对随机产生的伪负样本进行评估, 减小其落入适宜地区的概率, 进一步提高需要正负样本的II类模型的准确率。GIS功能在PSDS 2.0中也得到加强, 被应用于数据准备及结果分析等重要环节。文章最后以白冠长尾雉(Syrmaticus reevesii)为例, 运用PSDS 2.0系统预测其在中国范围内的潜在分布地, 并对各种模型的预测结果进行评估和比较。     

使用大熊猫数据评估Biomod2和MaxEnt分布预测模型的表现

物种分布模型是物种研究和保护者常用的工具.不同模型的预测结果可能相差很大,对研究者选择模型造成一定的难度.本研究使用大熊猫的实际分布数据评估了两种常见物种分布模型Biomod2和最大熵模型(MaxEnt)的表现,运用ROC曲线下面积(area under the curve,AUC)、真实技巧统计值(true skill statistics,TSS)、KAPPA统计量3种指标综合评估了两种模型预测结果的准确度.结果表明: 当使用的物种分布数据和模拟重复次数足够多的时候,两者都能够给出相当准确的预测.相对于MaxEnt,Biomod2的预测准确度更高,尤其是在物种分布点稀少的情况下.然而,Biomod2使用难度较大,运行时间较长,数据处理能力有限.研究者应基于对预测结果的误差要求来选择模型.在误差要求明确且两个模型都能满足误差要求时,建议使用MaxEnt,否则应优先考虑使用Biomod2.     

Modeling potential invasion range of alien invasive species,Hyptis suaveolens (L.) Poit. in India: Comparison of MaxEnt and GARP

Bushmint (Hyptis suaveolens (L.) Poit.) is one among the world's most noxious weeds. Bushmint is rapidly invading tropical ecosystems across the world, including India, and is major threat to native biodiversity, ecosystems and livelihoods. Knowledge about the likely areas under bushmint invasion has immense importance for taking rapid response and mitigation measures. In the present study, we model the potential invasion range of bushmint in India and investigate prediction capabilities of two popular species distribution models (SDM) viz., MaxEnt (Maximum Entropy) and GARP (Genetic Algorithm for Rule-Set Production). We compiled spatial layers on 22 climatic and non-climatic (soil type and land use land cover) environmental variables at India level and selected least correlated 14 predictor variables. 530 locations of bushmint along with 14 predictor variables were used to predict bushmint distribution using MaxEnt and GARP. We demonstrate the relative contribution of predictor variables and species-environmental linkages in modeling bushmint distribution. A receiver operating characteristic (ROC) curve was used to assess each model's performance and robustness. GARP had a relatively lower area under curve (AUC) score (AUC: 0.75), suggesting its lower ability in discriminating the suitable/unsuitable sites. Relative to GARP, MaxEnt performed better with an AUC value of 0.86. Overall the outputs of MaxEnt and GARP matched in terms of geographic regions predicted as suitable/unsuitable for bushmint in India, however, predictions were closer in the spatial extent in Central India and Western Himalayan foothills compared to North-East India, Chottanagpur and Vidhayans and Deccan Plateau in India.     

ROC曲线分析在评价入侵物种分布模型中的应用

生态位模型（ecological niche models,ENMs）已广泛应用于物种潜在分布区预测,ENMs的应用也为外来入侵物种的风险分析提供了重要的定量化分析工具,但如何评价不同模型之间的预测效果成了当今研究的热点问题。本文介绍了受试者工作特征（ROC）曲线分析在评价不同生态位模型预测效果中的应用原理和分析方法,并以一种植物病原线虫-相似穿孔线虫（Radopholus similis）为例,应用ROC曲线分析法对其5种模型（BIOCLIM,CLIMEX,DOMAIN,GARP,MAXENT）的预测结果进行了比较分析。5种模型的ROC曲线下面积AUC（Area Under Curve）值分别为0.810,0.758,0.921,0.903和0.950,以MAXENT模型的AUC值最大,表明其预测效果最好;方差分析结果表明,除GARP与DOMAIN模型之间AUC值差异不显著外,其余各模型之间差异显著。     

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.     

An evaluation of a GARP model as an approach to predicting the spatial distribution of non-vagile invertebrate species

One of the primary goals of any systematic, taxonomic or biodiversity study is the characterization of species distributions. While museum collection data are important for ascertaining distributional ranges, they are often biased or incomplete. The Genetic Algorithm for Rule-set Prediction (GARP) is an ecological niche modelling method based on a genetic algorithm that has been argued to provide an accurate assessment of the spatial distribution of organisms that have dispersal capabilities. The primary objective of this study is to evaluate the accuracy of a GARP model to predict the spatial distribution of a non-invasive, non-vagile invertebrate whose full distributional range was unknown. A GARP predictive model based on seven environmental parameters and 42 locations known from historical museum records for species of the trapdoor spider genus Promyrmekiaphila was produced and subsequently used as a guide for ground truthing the model. The GARP model was neither a significant nor an accurate predictor of spider localities and was outperformed by more simplistic BIOCLIM and GLM models. The isolated nature of Promyrmekiaphila populations mandates that environmental layers and their respective resolutions are carefully chosen for model production. Our results strongly indicate that, for modelling the spatial distribution of low vagility organisms, one should employ a modelling method whose results are more conducive to interpretation than models produced by a 'black box' algorithm such as GARP.     

利用最大熵模型和规则集遗传算法模型预测孑遗植物裸果木的潜在地理分布及格局

利用野外调查的16个居群分布点和7个环境因子图层, 选择最大熵模型(MAXENT)和规则集遗传算法模型(GARP), 在地理和环境空间上模拟了第三纪孑遗植物裸果木(Gymnocarpos przewalskii)在中国西北地区的潜在分布。结果表明: (1)裸果木的潜在适生区全部集中在西北荒漠区, 其中最佳适生区主要集中在3个区域, 一是河西走廊中部和玉门以西、宁夏北部及内蒙古乌拉特后旗; 二是塔里木盆地西北缘; 三是柴达木盆地西北缘两片极小的高度适生区。裸果木的生态位被确定在一个较广的干旱环境空间: 适生区极端最高气温基本上在29.2-36.8 ℃之间, 极端最低气温在-18.3至-13.4 ℃之间; 年平均降水量40-200 mm; 潜在蒸发率在3-15之间。(2) MAXENT和GARP模型都较好地预测了裸果木的潜在分布, 但GARP产生了相对较大、较连续的潜在分布区, 部分过预测了破碎化生境; 而MAXENT预测到的潜在分布区, 在不同区域具有不同的环境适生性指数, 而且成功地排除了不合理的破碎化分布, 从而更直观地展示了裸果木的潜在分布格局和生态位要求。     

The effect of sample size and species characteristics on performance of different species distribution modeling methods

Species distribution models should provide conservation practioners with estimates of the spatial distributions of species requiring attention. These species are often rare and have limited known occurrences, posing challenges for creating accurate species distribution models. We tested four modeling methods (Bioclim, Domain, GARP, and Maxent) across 18 species with different levels of ecological specialization using six different sample size treatments and three different evaluation measures. Our assessment revealed that Maxent was the most capable of the four modeling methods in producing useful results with sample sizes as small as 5, 10 and 25 occurrences. The other methods compensated reasonably well (Domain and GARP) to poorly (Bioclim) when presented with datasets of small sample sizes. We show that multiple evaluation measures are necessary to determine accuracy of models produced with presence-only data. Further, we found that accuracy of models is greater for species with small geographic ranges and limited environmental tolerance, ecological characteristics of many rare species. Our results indicate that reasonable models can be made for some rare species, a result that should encourage conservationists to add distribution modeling to their toolbox.     

样本容量和物种特征对BIOCLIM模型模拟物种分布准确度的影响——以12个中国特有落叶栎树种为例

物种分布模型被广泛应用于生态学、生物地理学及保护生物学等领域的研究。由于难于取样或标本记录不完善等原因, 真正能够用于模型预测的物种分布数据非常有限。因此, 有必要搞清楚样本容量和物种特征对模型模拟准确度的影响, 为确定以物种特征为区分条件的最小样本容量奠定基础。为了探讨应用BIOCLIM模型预测中国特有植物种的效果, 以12个落叶栎树种为例, 从不同的样本容量和生态特征两方面研究其对BIOCLIM模型模拟准确度的影响。结果表明: BIOCLIM模型模拟准确度随样本容量的增加在初期几乎呈直线增加趋势至样本容量达到25, 随后渐变平缓至样本容量为75~100时达到最大值。此外, 生态幅窄和环境特化物种比生态幅宽和对环境耐受性强的物种更容易获得较高的准确度。结果说明, BIOCLIM可有效地用于样本数量较小的狭域型物种分布预测。     

Predicting species distributions from small numbers of occurrence records: a test case using cryptic geckos in Madagascar

Aim Techniques that predict species potential distributions by combining observed occurrence records with environmental variables show much potential for application across a range of biogeographical analyses. Some of the most promising applications relate to species for which occurrence records are scarce, due to cryptic habits, locally restricted distributions or low sampling effort. However, the minimum sample sizes required to yield useful predictions remain difficult to determine. Here we developed and tested a novel jackknife validation approach to assess the ability to predict species occurrence when fewer than 25 occurrence records are available. Location Madagascar. Methods Models were developed and evaluated for 13 species of secretive leaf‐tailed geckos (Uroplatus spp.) that are endemic to Madagascar, for which available sample sizes range from 4 to 23 occurrence localities (at 1 km² grid resolution). Predictions were based on 20 environmental data layers and were generated using two modelling approaches: a method based on the principle of maximum entropy (Maxent) and a genetic algorithm (GARP). Results We found high success rates and statistical significance in jackknife tests with sample sizes as low as five when the Maxent model was applied. Results for GARP at very low sample sizes (less than c. 10) were less good. When sample sizes were experimentally reduced for those species with the most records, variability among predictions using different combinations of localities demonstrated that models were greatly influenced by exactly which observations were included. Main conclusions We emphasize that models developed using this approach with small sample sizes should be interpreted as identifying regions that have similar environmental conditions to where the species is known to occur, and not as predicting actual limits to the range of a species. The jackknife validation approach proposed here enables assessment of the predictive ability of models built using very small sample sizes, although use of this test with larger sample sizes may lead to overoptimistic estimates of predictive power. Our analyses demonstrate that geographical predictions developed from small numbers of occurrence records may be of great value, for example in targeting field surveys to accelerate the discovery of unknown populations and species.     

ROC Curve Analysis in the Presence of Imperfect Reference Standards

The receiver operating characteristic (ROC) curve is an important tool for the evaluation and comparison of predictive models when the outcome is binary. If the class membership of the outcomes is known, ROC can be constructed for a model, and the ROC with greater area under the curve indicates better performance. However in practice, imperfect reference standards often exist, in which class membership of every data point is not fully determined. This situation is especially prevalent in high-throughput biomedical data because obtaining perfect reference standards for all data points is either too costly or technically impractical. To construct ROC curves for these data, the common practice is to either ignore the uncertainties in references or remove data points with high uncertainties. Such approaches may cause bias to the ROC curves and generate misleading results in method evaluation. Here we present a framework to incorporate membership uncertainties into the construction of ROC curve, termed the expected ROC or “eROC” curve. We develop an efficient procedure for the estimation of eROC curve. The advantages of using eROC are demonstrated using simulated and real data.     

基于GARP的三种芒果象甲在中国的适生性分析

芒果象甲Sternochetus Pierce昆虫是芒果的重要害虫,明确其可能适牛的区域对该虫的科学监测及防治意义重大.本文利用芒果象甲属中主要3种:印度果核芒果象S.mangiferae Fabricius、果核芒果象S.olivieri Faust、果肉芒果象S.frigidus Fabrieius的已知分布点数据和亚洲地区的14个环境地理变量图层,运用GARP生态位模型结合GIS空间分析模块预测了该虫在中国的潜在地理分布,结果表明芒果象甲具有较强的扩散蔓延趋势,对我国的芒果产业构成较大的潜在威胁.