北美水貂和欧亚水獭在东北地区的分布与生态位重叠
Distribution and niche overlap of American mink and Eurasian otter in Northeast China
通讯作者: *E-mail:luanxiaofeng@bjfu.edu.cn
编委: 乔慧捷
责任编辑: 黄祥忠
收稿日期: 2022-05-27 接受日期: 2022-08-31
基金资助: |
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Corresponding authors: *E-mail:luanxiaofeng@bjfu.edu.cn
Received: 2022-05-27 Accepted: 2022-08-31
作为入侵物种, 北美水貂(Neovison vison)在欧洲引起了一系列生态问题, 侵占了欧亚水獭(Lutra lutra)的生态空间, 其入侵性对当地生物多样性和生态系统构成了严重威胁。水貂引入我国东北地区已有70多年的历史, 然而国内对其野外种群却鲜有研究。掌握水貂种群的入侵范围、入侵影响因素以及与本地具有相似生态位的欧亚水獭之间的竞争关系, 对水貂的入侵管理和东北地区的生物多样性保护具有重要意义。本研究利用实地调查和文献资料获取的分布信息, 通过集合模型识别水貂和水獭的潜在分布区, 评估水貂对水獭在地理空间上的入侵风险, 并通过主成分分析(principal component analysis, PCA)评估其生态位重叠和影响因素。结果表明: (1)我国东北地区水貂的潜在分布区面积为61,944.57 km2, 水獭的潜在分布区面积为83,590.94 km2, 两者重叠区域面积为50,544.21 km2, 占水獭潜在分布区面积的60.47%; (2)从各省分布情况来看, 黑龙江省水獭受水貂入侵的风险最高, 潜在分布区重叠的比例达到78.94%, 其次是吉林省, 重叠比例约为53.80%; (3)水貂和水獭生态位存在高度重叠, Schoener’s D值达到0.60。单因子生态位分析结果表明, 水獭比水貂对耕地密度更加敏感, 集中分布在低耕地密度区; 水獭和水貂均倾向选择有林地密度较高的区域, 但水貂在有林地密度的选择上更加平滑, 对森林的依赖程度比水獭低; 水貂相比水獭更倾向选择降水量低的区域; 水獭比水貂更倾向选择高海拔的栖息地。根据地理分布特征和生态位分析的结果, 我们建议通过控制人为干扰调控水獭和水貂的竞争关系, 以抑制水貂在东北地区的入侵。
关键词:
Aims: As an invasive species, the American mink (Neovison vison) has caused a series of ecological problems in Europe. It competes for ecological space with the Eurasian otter (Lutra lutra), and its invasiveness poses a serious threat to local biodiversity and ecosystem. However, minks have been introduced into the Northeast China for more than 70 years, yet there are few studies on the field population of minks in China. The purpose of this study is to understand the influencing factors and invasive range distribution of the mink population and the competition relationship between minks and otters with a similar ecological niche.
Methods: In this study, based on the distribution information obtained from field investigation and literature, the potential distribution areas of American mink and Eurasian otter were identified by utilizing an ensemble model, and the geospatial invasion risk of American mink to Eurasian otter was assessed. Principal component analysis (PCA) was used to assess niche overlap and influencing factors.
Results: The potential distribution area of American mink was 61,944.57 km2, while the potential distribution area of Eurasian otter was 83,590.94 km2, and the overlapping area was 50,544.21 km2, accounting for 60.47% of the potential distribution area of otters in Northeast China. With distribution by province, otters in Heilongjiang Province had the highest risk of invasion by minks, and the overlap ratio of potential distribution area is 78.94%, followed by Jilin province with an overlap of 53.80%. The environmental niche of minks and otters was highly overlapped, with Schoener’s D value reaching 0.60. The results of single-factor niche analysis indicated that otters were more sensitive to cultivated land density than minks and concentrate in low cultivated land density areas. Otters and minks tended to choose areas with higher forest density, but minks had a smoother selection of forest density and were less dependent on forest than otters. Minks preferred areas with lower precipitation than otters and otters preferred higher-altitude habitats to minks.
Conclusion: According to the geographical distribution characteristics and the results of niche analysis, we suggest that the competition between otter and mink should be regulated by controlling human disturbance to suppress the mink invasion in Northeast China.
Keywords:
本文引用格式
陈敏豪, 张超, 王嘉栋, 湛振杰, 陈君帜, 栾晓峰 (2023)
Minhao Chen, Chao Zhang, Jiadong Wang, Zhenjie Zhan, Junzhi Chen, Xiaofeng Luan (2023)
全球化进程让各国间的商品贸易和旅游活动更加频繁, 高频次的人员和物资交流为生物入侵提供了便利。自19世纪以来, 入侵物种的数量迅速增加, 尤其是肉食性哺乳类, 通过捕食和竞争对本土物种造成严重威胁(Courchamp et al, 2003; Gurevitch & Padilla, 2004; Seebens et al, 2017)。目前, 生物入侵已被认为是全球生物多样性丧失的主要驱动因素之一(Sala et al, 2000; Perrings et al, 2010; Keller et al, 2011)。入侵物种和本土物种的相互作用是了解生物入侵进程的重要前提, 可为识别和预测潜在的入侵区域、制定防控策略提供基本信息(Qiao et al, 2017)。在生物入侵中, 入侵物种与本土具相似生态位的物种会发生竞争(Bøhn et al, 2008; Almeida Saá et al, 2020)。如果这种竞争关系在整个范围内对本土物种丰度产生持续的负面影响, 则本土物种可能会因竞争排斥而减少甚至灭绝(Bøhn et al, 2008; Bell et al, 2021)。种间竞争的相对程度可以通过生态位重叠来测度(Abrams, 1980; Broennimann et al, 2012), 从而在一定程度上量化入侵物种对本地相似生态位物种的影响, 以支持入侵物种的政策决策和管理工作(Qiao et al, 2017; Alves et al, 2021)。
在入侵性哺乳动物中, 北美水貂(Neovison vison)已经造成了各种各样的保护问题, 它们建立了广泛的野生种群, 对本土物种, 特别是鸟类、两栖类、小型啮齿类和鼬科动物产生了显著影响(Banks et al, 2004; Nordström & Korpimäki, 2004; Bonesi & Palazon, 2007)。水貂作为重要经济毛皮动物在20世纪中叶引入我国东北地区, 并在黑龙江和吉林建立起野生种群(郭文场和杨智奎, 1963), 通过对野生和饲养的水貂下颌骨形态特征差异的分析, 也证实了水貂在东北地区野外环境下的适应性进化和野外稳定种群的建立(刘博洋等, 2021)。东北地区还是我国欧亚水獭(Lutra lutra)的重要分布区之一(Zhang et al, 2018; 张超等, 2022)。研究表明, 水貂和水獭存在食物和栖息空间上的竞争关系, 且水獭食性比水貂更窄。水貂以甲壳类、鱼类、小型哺乳类和鸟类为食, 其中甲壳类占较大比例; 水獭则主要以鱼类为食, 甲壳类是其次要食物来源(Clode & MacDonald, 1995)。两者均栖息于河流和湖泊岸带, 尤其喜欢生活在两岸林木繁茂的溪河地带。在过去70多年间, 东北地区的水獭因盗猎、栖息地破碎化和水质污染等多重因素的影响数量下降了90%以上(朴正吉等, 2011; Zhang et al, 2016), 为水貂的入侵提供了契机, 导致空置的生态位被迅速占领, 水貂种群在过去几十年间迅速扩大(朴正吉等, 2011; 罗玉梅等, 2012)。
生物入侵是抑制种群恢复的重要因素之一。近年来调查显示, 东北地区的水獭种群正在恢复, 但该区域水貂的入侵范围和程度目前仍不清楚, 了解水貂和水獭间的生态位重叠关系, 量化入侵风险, 将为水獭保护和区域入侵种防治提供理论参考。本研究基于2016-2021年的调查和近30年的文献资料记录获取水貂和水獭的分布信息, 利用R软件中的BIOMOD2和ecospat程序包(Thuiller et al, 2009; Di Cola et al, 2017), 结合GIS工具, 通过集合模型模拟东北地区水貂和水獭的潜在分布区, 并利用排序技术构建其生态位。在此基础上, 探讨了东北地区水貂的分布格局和入侵风险, 分析了水貂与水獭间的生态位重叠程度和重要影响因子。
1 材料与方法
1.1 数据收集
图1
图1
中国东北地区北美水貂和欧亚水獭分布点
Fig. 1
Distribution sites of American mink and Eurasian otter in Northeast China
根据物种的生理特征和栖息地需求(Previtali et al, 1998; Robitaille & Laurence, 2002; Zabala et al, 2007; 张超等, 2022), 我们选取了16个环境变量, 包括: (1) 3个气候变量, 即年降水量、最湿季降水量、最干季降水量; (2) 4个地形变量, 即海拔、坡向、坡度和地表起伏度; (3) 5个地类变量, 即距道路距离、耕地密度、有林地密度、居民点密度和距城镇距离; (4) 4个与水域有关的变量, 即距水域距离、距主要河流距离、距一级支流距离、距二级支流距离。其中, 坡向和坡度由海拔数据生成; 耕地密度、有林地密度、居民点密度和距城镇距离由土地利用类型数据提取相应地类栅格, 利用ArcGIS的核密度分析和欧氏距离分析程序, 按默认参数计算并进行归一化处理得到。由于水貂和水獭的分布与河流大小也有关联, 我们提取了多年平均径流量大于1亿m3的河流作为主要河流(附录1), 并以此划分一级支流和二级支流, 分别生成距离图层。环境变量分辨率统一定义为30″ (约1 km)。
1.2 物种分布模拟
在物种分布模型(species distribution model, SDM)的研究中, 相比单一模型, 集合模型在模拟精度和可视化效果方面表现较好。因此, 我们选用基于R软件开发的BIOMOD2程序包进行物种分布模拟。BIOMOD2包含10种模型, 分别是广义线性模型(GLM)、广义加法模型(GAM)、广义推进模型(GBM)、分类树模型(CTA)、人工神经网络模型(ANN)、柔性判别分析(FDA)、表面分布区分室模型(SRE/BIOCLIM)、多元适应回归样条函数(MARS)、随机森林(RF)和最大熵(MaxEnt)模型(Thuiller et al, 2009)。环境因子的多重共线性会影响物种分布模型的准确性, 为此, 我们通过方差膨胀因子(variance inflation factor, VIF)的两次迭代运算筛选出15个环境变量(表1), 以降低变量间的过度拟合(VIF < 10, 附录2), 并利用dismo程序包在研究区内随机生成5,000个伪缺失点用于模型构建。为了降低模型的不确定性, 我们选取80%的数据作为训练样本, 20%的数据作为验证样本, 经过10次重复运算, 选择平均真实技巧统计值(true skill statistics, TSS)和平均受试者操作特征曲线下的面积值(area under curve, AUC)最高的5个模型进行30次重复运算集合建模(附录3)。最终, 基于模型评价分数计算的阈值(cut-off)将研究区域划分为适宜分布区和不适宜分布区。
表1 环境变量的重要性及建模选择。年降水量在使用方差膨胀因子评估后被排除; ● 被选择的环境变量; × 被排除的环境变量。
Table 1
环境变量 Environmental variables | 重要性 Importance | 平方和的平方根 RSS | 物种分布 模型 SDM | 生态位量化 Niche quantification* | 数据来源 Data sources | |
---|---|---|---|---|---|---|
水貂 Mink | 水獭 Otter | |||||
距水域距离 Distance from water | 0.765 | 0.692 | 1.031 | ● | ● | 全国地理信息资源目录服务系统 National Catalogue Service for Geographic Information https://www.webmap.cn/ |
居民点密度 Residential density | 0.124 | 0.036 | 0.129 | ● | ● | 资源环境科学与数据中心 Resource and Environment Science and Data Center http://www.resdc.cn/Default.aspx/ |
耕地密度 Cultivated land density | 0.037 | 0.112 | 0.118 | ● | ● | 资源环境科学与数据中心 Resource and Environment Science and Data Center http://www.resdc.cn/Default.aspx/ |
最干季降水量 Precipitation of driest quarter | 0.056 | 0.056 | 0.080 | ● | ● | 世界气候数据网站 WorldClim http://www.worldclim.org/ |
海拔 Elevation | 0.070 | 0.005 | 0.071 | ● | ● | SRTM 90 m数字高程数据 SRTM 90 m Digital Elevation Database http://srtm.csi.cgiar.org/ |
有林地密度 Forest density | 0.021 | 0.029 | 0.036 | ● | ● | 资源环境科学与数据中心 Resource and Environment Science and Data Center http://www.resdc.cn/Default.aspx/ |
最湿季降水量 Precipitation of wettest quarter | 0.014 | 0.001 | 0.014 | ● | ● | 世界气候数据网站 WorldClim http://www.worldclim.org/ |
坡度 Slope | 0.008 | 0.002 | 0.009 | ● | × | SRTM 90 m数字高程数据 SRTM 90 m Digital Elevation Database http://srtm.csi.cgiar.org/ |
地表起伏度 Relief degree of land surface | 0.007 | 0.004 | 0.008 | ● | × | 全球变化科学研究数据出版系统 Global Change Research Data Publishing and Repository http://www.geodoi.ac.cn/ |
距一级支流距离 Distance from primary tributaries | 0.004 | 0.007 | 0.008 | ● | × | 全国地理信息资源目录服务系统 National Catalogue Service for Geographic Information https://www.webmap.cn |
距二级支流距离 Distance from secondary tributaries | 0.007 | 0.002 | 0.008 | ● | × | 全国地理信息资源目录服务系统 National Catalogue Service for Geographic Information https://www.webmap.cn |
坡向 Aspect | 0.005 | 0.003 | 0.006 | ● | × | SRTM 90 m数字高程数据 SRTM 90 m Digital Elevation Database http://srtm.csi.cgiar.org/ |
距主要河流距离 Distance from major rivers | 0.004 | 0.001 | 0.004 | ● | × | 全国地理信息资源目录服务系统 National Catalogue Service for Geographic Information https://www.webmap.cn |
距道路距离 Distance from road | 0.002 | 0.000 | 0.002 | ● | × | 全国地理信息资源目录服务系统 National Catalogue Service for Geographic Information https://www.webmap.cn |
距城镇距离 Distance from town | 0.001 | 0.001 | 0.001 | ● | × | 资源环境科学与数据中心 Resource and Environment Science and Data Center http://www.resdc.cn/Default.aspx |
年降水量 Annual precipitation | - | - | - | × | × | 世界气候数据网站 WorldClim http://www.worldclim.org/ |
* 生态位量化分析中逐一去除平方和的平方根较小的变量, 当剩余前7个环境变量时, 主成分分析(PCA)结果中PC3特征值小于1, 因此使用前7个环境变量进行生态位量化分析。
* In niche quantification analysis, variables with smaller root sum square (RSS) were removed one by one. When the first 7 environmental variables were left, the eigenvalue of PC3 in principal component analysis (PCA) results was less than 1, so the first 7 environmental variables were used for niche quantification analysis. SDM, Species distribution model.
1.3 生态位重叠测量
排序技术中的主成分分析(principal component analysis, PCA)被广泛运用于物种生态位分析中(Di Cola et al, 2017)。在测量生态位重叠时, 与物种分布模型不同, 排序技术允许直接比较空间中的物种和环境的关系, 采用各种最大化标准从相关环境变量构建合成轴, 并且不会因环境梯度的范围和分布而变化(Jongman et al, 1995; Broennimann et al, 2012)。因此, 我们选用R软件中基于PCA开发的ecospat程序包进行生态位量化分析(Thuiller et al, 2009)。为了排除统计学上与两个物种分布相关性较低的环境变量, 并提高PCA中前两个成分的方差贡献率, 根据两个物种分布模拟中环境变量重要性平方和的平方根(root sum square, RSS, 表1), 逐个去掉排在后面的变量进行PCA。当PC1和PC2的累计方差贡献率大于85%或PC3特征值小于1时, 选择剩余的变量(当执行至剩余前7个变量时, PC3特征值为0.832, 满足条件)构建其生态位(覃林, 2009)。由于生态位分析和物种分布模型对分布数据的要求不同(Di Cola et al, 2017), 我们先使用未稀疏化的所有560个出现点和5,000个伪缺失点作为环境背景值对PCA进行校准, 再将两个物种分布的PCA分数投射到研究区域内以最小和最大PCA分数为边界的空间中, 利用核密度函数使用100个像素单位进行搜索计算, 核密度平滑后计算Schoener’s D值(Schoener, 1970), 用以衡量生态位重叠程度, 并对重要因子生态位特征进行分析。其中, Schoener’s D取值范围0-1, 0表示完全没有重叠, 1表示完全重叠。
2 结果
2.1 生态位模型选择与模型表现
在两个物种的分布模拟中, 模型预测均达到较高精度。在水貂的分布模拟中, 采用GAM、GBM、GLM、RF和MaxEnt进行组合建模, 平均TSS值达0.923, 平均AUC值达0.992; 在水獭的分布模拟中, 选择GAM、GBM、GLM、RF和MARS组合建模, 平均TSS值达0.918, 平均AUC值达0.994。
2.2 环境因子重要性
根据物种分布模型的评估结果, 影响水貂分布最重要的环境因子是距水域距离、居民点密度和海拔; 影响水獭分布最重要的环境因子是距水域距离、耕地密度和最干季降水量。根据环境变量重要性的结果(表1), 我们排除了与两个物种分布相关性较低的环境变量, 保留距水域距离、居民点密度、耕地密度、最干季降水量、海拔、有林地密度和最湿季降水量共7个环境变量构建水貂和水獭生态位。
2.3 潜在分布区及对水獭的入侵风险
表2 东北地区北美水貂和欧亚水獭潜在分布区及重叠的比例
Table 2
省份 Province | 潜在分布区 Potential distribution (km2) | 重叠比例 Proportion of overlap | ||
---|---|---|---|---|
水貂 Mink | 水獭 Otter | 重叠区域 Overlapping region | ||
内蒙古 Inner Mongolia | 12,616.02 | 27,758.67 | 12,135.05 | 43.72% |
辽宁 Liaoning | 283.56 | 2,780.04 | 266.35 | 9.58% |
吉林 Jilin | 8,264.06 | 14,871.51 | 8,001.23 | 53.80% |
黑龙江 Heilongjiang | 40,780.93 | 38,180.71 | 30,141.58 | 78.94% |
总计 Total | 61,944.57 | 83,590.94 | 50,544.21 | 60.47% |
图2
图2
东北地区北美水貂和欧亚水獭的潜在分布区
Fig. 2
Potential distribution areas of American mink and Eurasian otter in Northeast China
2.4 生态位分析
在物种分布模型的基础上, 利用PCA构建和测量水貂和水獭的生态位重叠程度。基于所有环境背景值的PCA结果显示, PC1和PC2分别解释了整体变异性的35.71%和31.63%, 共67.34%。PC1主要与有林地密度和耕地密度相关, PC2主要与最湿季降水量和海拔相关。其中, 耕地密度对结果解释性最高,其次是有林地密度和最湿季降水量(图3)。结合物种生态位重叠分析的结果, 水貂生态位沿PC1方向扩张, 但种间生态位密度核心与水獭存在高度重叠, Schoener’s D值达到0.60; 水獭生态位在沿PC2方向上与水貂存在较大的未重叠区域, 表明两者在最湿季降水量和海拔的生态位占据上存在较大差异。
图3
图3
基于主成分分析(PCA)的生态位重叠分析结果。
A: 所有环境背景值的主成分分析相关性, cos2值表示各个因子对结果的解释性, 高cos2值表示该变量在主成分上具有良好的解释性, 且指向终点越靠近圆周。B: 水貂和水獭的生态位重叠, 实线和虚线轮廓线分别显示了可用背景环境的100%和50%, 绿色为水獭占据的生态位, 红色为水貂占据的生态位, 紫色为两个物种生态位的重叠部分, Schoener’s D值为0.60。C-F: 单因子生态位分布特征,实线和虚线轮廓线分别显示了可用背景环境的100%和50%, 绿色为水獭占据的生态位, 红色为水貂占据的生态位, 紫色为两个物种生态位的重叠部分; 横坐标轴上方的色条中, 绿色为背景环境的50%中被水獭单独占据的区域, 红色为背景环境的50%中被水貂单独占据的区域, 蓝色为背景环境的50%中共同占据的区域。
Fig. 3
Niche overlap analysis results based on principal component analysis (PCA).
A, Correlation circle of PCA of all environmental background values, cos2 value indicates the interpretation of each factor to the results, high cos2 value indicates that the variable has good interpretation on the principal component, and points closer to the circumference of the end point. B, The niche overlap of mink and otter. The solid and dotted lines show 100% and 50% of the available background environment, respectively. The green is the ecological niche occupied by otters, the red is the ecological niche occupied by minks, and the purple is the overlapped niche of the two species, with Schoener’s D value of 0.60. C-F, Single factor niche distribution characteristics, solid line and dotted line contour lines show 100% and 50% of available background environment, respectively, the green is the niche occupied by otters, the red is the niche occupied by minks, and the purple is the overlapping niche of the two species. In the color bar above the horizontal axis, the green is the area occupied by otter alone in 50% of the background environment, the red is the area occupied by mink alone in 50% of the background environment, and the blue is the area occupied jointly in 50% of the background environment.
单因子生态位分析结果表明, 水獭比水貂对耕地密度更加敏感, 分布密度峰值偏离背景环境密度峰值, 集中分布在低耕地密度区。在背景环境生态位存在低密度和高密度两个峰的情况下, 水獭和水貂均倾向选择有林地密度较高的区域, 且在水獭种群内出现两个不同的分布峰值, 而水貂的分布密度在有林地密度的选择上更加平滑, 对森林的依赖程度比水獭低。最湿季降水量与区域集水和河流大小有关, 尽管水貂和水獭在该因子的生态位分布中高度重叠, 但水貂分布密度峰值偏离背景环境密度峰值, 更倾向选择降水量低的区域。水貂和水獭的分布在海拔上趋向选择比可用背景环境海拔峰值更高的区域, 但水獭表现得更加明显, 分布密度峰值偏离背景环境密度峰值和水貂的分布峰值。
3 讨论
3.1 物种分布模拟
本研究结合多来源分布数据交叉验证和集合模型, 在模拟中增加与河流规模相关的环境变量, 提高了识别和预测水貂和水獭潜在分布区的准确性。物种分布数据带来的不确定性会影响对模型结果的解释, 并降低其在保护规划中的有效性(Tang et al, 2020)。我们通过访问调查、问卷调查、野外实地调查、粪便DNA鉴定和文献回顾等多种方式对水貂和水獭的分布点进行收集和交叉验证, 在以往研究的基础上, 补充了长白山地区以及辽宁部分分布点(吕江等, 2018; 张超等, 2022)。除气候、地形和植被因素外, 水貂和水獭栖息地的选择与环境污染和河流规模有关, 因而我们在模拟中增加了与农业污染相关的耕地密度以及河流等级。在识别和预测潜在入侵区域时, 单一的物种分布模型结果往往是多变和不可靠的, 而集合模型的方法可以提高模型精度, 降低模型的不确定性(Guo et al, 2015)。本研究通过模型测试选择精度较高的多个算法进行集合建模, 增加了模拟结果的可靠性。
3.2 生态位量化
量化物种分布并确定哪些因素影响物种分布范围一直以来都是生态学研究的主题之一(Guisan & Thuiller, 2005)。得益于生态位概念的发展(Grinnell, 1917), 许多方法和指标被用于量化物种生态位特征和不同物种的生态位重叠, 尤其是排序技术和物种分布模型应用广泛(Guisan & Thuiller, 2005; Thuiller et al, 2005)。排序技术中的主成分分析能将多个相关变量转换为两个不相关线性组合, 使研究区域的生态变异最大化, 以排除环境梯度的影响, 从而在背景环境空间中比较物种占据的生态位(Di Cola et al, 2017)。针对高维稀疏特征或高维不相关的特征, 主成分分析存在信息丢失的风险。因此, 评估物种生态位的差异需要识别和考虑影响物种分布的关键因素, 以降低环境变量维度。物种分布模型可以通过加权选择强调决定物种分布相关的某些变量, 排除不利于区分物种存在与否的变量或降低其权重, 并在地理空间中投影呈现(Wintle et al, 2003; Guisan & Thuiller, 2005)。结合物种分布模型中环境因子重要性的排序结果, 选择影响物种分布的关键变量用于生态位量化, 可降低环境变量维度, 从而提高PC1和PC2的累计方差贡献率, 以规避信息丢失。
3.3 竞争共存和管控建议
模型模拟的结果表明, 水貂潜在的入侵范围主要集中在大兴安岭靠近黑龙江的区域、小兴安岭腹地、三江平原和长白山地区, 除三江平原外, 在其他区域均与水獭形成分布上的高度重叠(图2)。已有证据证明水獭在与水貂的食物竞争中处于优势, 能在一定程度上抑制水貂种群的发展, 当生境中存在除鱼类和甲壳类之外的其他猎物时, 水貂为了避免与水獭竞争而增加其他猎物的捕食比例, 但同时出现身体状况的下降(Harrington et al, 2009)。尽管水獭捕获水生猎物的能力优于水貂, 但水貂高度的生态适应能力也使其在复杂的竞争环境中通过改变饮食结构与水獭实现共存(Clode & MacDonald, 1995; Harrington et al, 2009)。在两者竞争过程中, 物种对人为干扰的耐受程度也发挥了重要作用, 除了人为猎杀和栖息地破碎化, 有机氯等农业污染也是导致水獭种群急剧下降的重要因素(Chanin & Jefferies, 1978; Jefferies, 1989), 而水貂则在许多研究中被作为水生环境污染程度的哨兵物种(sentinel species), 对环境污染有一定的耐受性(Basu et al, 2007)。本研究结果表明, 在人为干扰的影响下, 两者的竞争可能存在不同的结果。在低海拔、耕地密度大的三江平原, 水貂具有更强的适应能力; 高海拔、耕地密度较低的大兴安岭主脉、长白山脉部分地区处于河流上游, 污染少, 水獭占据了大部分栖息地; 而在中海拔的黑龙江大兴安岭地区、小兴安岭和长白山部分区域, 在人为干扰和食物竞争的共同影响下, 形成了水貂和水獭共存的局面(图2, 3)。
水貂作为外来物种对本土物种的影响是多方面的, 包括捕食、竞争和外来物种携带的动物源性传染病等(Zhang et al, 2022)。水貂存在过量捕杀猎物的行为, 甚至1只水貂就可能会使整个栖息地中地面筑巢的鸟类数量锐减(Aars et al, 2001; Nordström & Korpimäki, 2004)。在发现水貂野生种群至今的70多年间, 其入侵导致的影响没有受到重视, 种群迅速繁衍导致部分区域的鱼类、两栖类和鸟类种群数量迅速下降(罗玉梅等, 2012)。有研究指出, 20世纪末欧洲水貂数量的下降可能与水獭种群的重建有关(Bonesi & MacDonald, 2004), 尽管可能不会直接导致水貂数量的长期显著减少, 但存在一定的抑制作用(García et al, 2009; Harrington et al, 2009)。因此, 可以利用水獭的竞争优势通过竞争置换的方式减少和控制水貂种群数量(Russell et al, 2014)。生态位分析的结果表明, 二者存在高度重叠的生态位, 但对耕地密度和海拔的生态位占据存在差异(图3)。结合两者的地理分布特征和生态位分析结果, 我们建议着重在中低海拔、耕地密度大的区域通过控制农药、化肥的使用和其他污染物以提高河流水质, 为水獭种群的恢复提供基本条件, 从而提高水獭在栖息地竞争中的优势以抑制水貂的数量; 在高海拔、人为干扰少的区域提高科研监测的投入, 监测并通过必要的捕杀以减少和控制水貂入侵的范围和影响。最新的研究表明, 外来物种携带的动物源性传染病也是对本土生物多样性和社会公共卫生健康的重要威胁之一(Swancutt & Chorba, 2021; Zhang et al, 2022)。水貂携带的犬瘟热病毒(canine distemper virus, CDV)会对当地野生食肉动物种群的生存能力构成严重威胁, 其余部分病毒也会对人类造成严重危害(Sepúlveda et al, 2014; Hammer et al, 2021)。因此, 加强对水貂饲养种群的逃逸防范管理和疾病控制, 对野外食肉动物, 尤其是对鼬科物种的保护十分重要。
3.4 不足之处
附录 Supplementary Material
附录1 东北地区主要河流多年平均径流量
Appendix 1 Mean annual runoff of major rivers in Northeast China
附录2 环境变量的方差膨胀因子(VIF)
Appendix 2 Variance inflation factor (VIF) value of environmental variables
附录3 使用真实技巧统计值(TSS)和受试者操作特征曲线下的面积值(AUC)选择最优建模算法
Appendix 3 The optimal modeling algorithm selected by using true skill statistics (TSS) and area under the receiver operating characteristic curve (AUC)
致谢
感谢内蒙古森工集团、呼伦贝尔市林草局、黑龙江省林草局、大兴安岭林业集团、黑河林草局、伊春森工集团、佳木斯林草局、吉林省林草局等单位对野外调查工作的大力支持,感谢长白山科学研究院的朴正吉和史国强、中山大学生命科学学院的杨立在数据分析和论文撰写等方面给予的帮助。
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DOI:10.1017/S1367943002004109 URL [本文引用: 1]
Over-invasion by functionally equivalent invasive species
Multiple invasive species have now established at most locations around the world, and the rate of new species invasions and records of new invasive species continue to grow. Multiple invasive species interact in complex and unpredictable ways, altering their invasion success and impacts on biodiversity. Incumbent invasive species can be replaced by functionally similar invading species through competitive processes; however the generalized circumstances leading to such competitive displacement have not been well investigated. The likelihood of competitive displacement is a function of the incumbent advantage of the resident invasive species and the propagule pressure of the colonizing invasive species. We modeled interactions between populations of two functionally similar invasive species and indicated the circumstances under which dominance can be through propagule pressure and incumbent advantage. Under certain circumstances, a normally subordinate species can be incumbent and reject a colonizing dominant species, or successfully colonize in competition with a dominant species during simultaneous invasion. Our theoretical results are supported by empirical studies of the invasion of islands by three invasive Rattus species. Competitive displacement is prominent in invasive rats and explains the replacement of R. exulans on islands subsequently invaded by European populations of R. rattus and R. norvegicus. These competition outcomes between invasive species can be found in a broad range of taxa and biomes, and are likely to become more common. Conservation management must consider that removing an incumbent invasive species may facilitate invasion by another invasive species. Under very restricted circumstances of dominant competitive ability but lesser impact, competitive displacement may provide a novel method of biological control.
Global biodiversity scenarios for the year 2100
DOI:10.1126/science.287.5459.1770
PMID:10710299
[本文引用: 1]
Scenarios of changes in biodiversity for the year 2100 can now be developed based on scenarios of changes in atmospheric carbon dioxide, climate, vegetation, and land use and the known sensitivity of biodiversity to these changes. This study identified a ranking of the importance of drivers of change, a ranking of the biomes with respect to expected changes, and the major sources of uncertainties. For terrestrial ecosystems, land-use change probably will have the largest effect, followed by climate change, nitrogen deposition, biotic exchange, and elevated carbon dioxide concentration. For freshwater ecosystems, biotic exchange is much more important. Mediterranean climate and grassland ecosystems likely will experience the greatest proportional change in biodiversity because of the substantial influence of all drivers of biodiversity change. Northern temperate ecosystems are estimated to experience the least biodiversity change because major land-use change has already occurred. Plausible changes in biodiversity in other biomes depend on interactions among the causes of biodiversity change. These interactions represent one of the largest uncertainties in projections of future biodiversity change.
Nonsynchronous spatial overlap of lizards in patchy habitats
DOI:10.2307/1935376 URL [本文引用: 1]
No saturation in the accumulation of alien species worldwide
DOI:10.1038/ncomms14435
PMID:28198420
[本文引用: 1]
Although research on human-mediated exchanges of species has substantially intensified during the last centuries, we know surprisingly little about temporal dynamics of alien species accumulations across regions and taxa. Using a novel database of 45,813 first records of 16,926 established alien species, we show that the annual rate of first records worldwide has increased during the last 200 years, with 37% of all first records reported most recently (1970-2014). Inter-continental and inter-taxonomic variation can be largely attributed to the diaspora of European settlers in the nineteenth century and to the acceleration in trade in the twentieth century. For all taxonomic groups, the increase in numbers of alien species does not show any sign of saturation and most taxa even show increases in the rate of first records over time. This highlights that past efforts to mitigate invasions have not been effective enough to keep up with increasing globalization.
Invasive American mink: Linking pathogen risk between domestic and endangered carnivores
DOI:10.1007/s10393-014-0917-z
PMID:24604545
[本文引用: 1]
Infectious diseases, in particular canine distemper virus (CDV), are an important threat to the viability of wild carnivore populations. CDV is thought to be transmitted by direct contact between individuals; therefore, the study of species interactions plays a pivotal role in understanding CDV transmission dynamics. However, CDV often appears to move between populations that are ecologically isolated, possibly through bridge hosts that interact with both species. This study investigated how an introduced species could alter multihost interactions and act as a bridge host in a novel carnivore assemblage of domestic dogs (Canis familiaris), invasive American mink (Neovison vison), and threatened river otters (Lontra provocax) in southern Chile. We found that rural dogs interact with mink near farms whereas in riparian habitats, minks and river otters shared the same latrines with both species visiting sites frequently within time intervals well within CDV environmental persistence. No interactions were observed between dogs and otters at either location. Both dog and mink populations were serologically positive for CDV, making the pathogen transfer risk to otters a conservation concern. Altogether, introduced mink in this ecosystem have the potential to act as bridge hosts between domestic dogs and endangered carnivores.
SARS-CoV-2, mannerism, marten, mink, and man
DOI:10.3201/eid2707.AC2707 URL [本文引用: 1]
Expanding ensembles of species present-day and future climatic suitability to consider the limitations of species occurrence data
Niche properties and geographical extent as predictors of species sensitivity to climate change
DOI:10.1111/j.1466-822X.2005.00162.x URL [本文引用: 1]
BIOMOD—A platform for ensemble forecasting of species distributions
DOI:10.1111/j.1600-0587.2008.05742.x URL [本文引用: 3]
The use of Bayesian model averaging to better represent uncertainty in ecological models
DOI:10.1111/j.1523-1739.2003.00614.x URL [本文引用: 1]
Winter habitat preferences of feral American mink Mustela vison in Biscay, Northern Iberian Peninsula
DOI:10.1007/BF03194196 URL [本文引用: 1]
Distribution pattern and identification of conservation priority areas of the otter in Northeast China
DOI:10.17520/biods.2021157
[本文引用: 3]
<p id="p00005"><strong>Aims:</strong> The otter is an indicator and flagship species of aquatic ecosystems. Its populations have undergone a drastic decline in China, and have become locally extinct in some regions due to anthropogenic disturbance. However, the current literature on otters in China is inadequate, which has subsequently affected conservation of the species. We aim to assess the potential distribution and conservation priority areas of the Eurasian otter (<i>Lutra lutra</i>) in Northeast China.</p><p id="p00010"><strong>Methods:</strong> In this study, we integrated species distribution models with otter survey data from 2016 to 2020 to assess Eurasian otter distribution. Then, we used the geographic information system and core-area zonation algorithm in Zonation 4.00 to identify conservation priority areas for otter protection. Using the anthropogenic pressures layer, we also evaluated anthropogenic stressors in each province. Then, we analyzed the conservation status of the otter based on the spatial distribution of national nature reserves. Furthermore, we used Inner Mongolia Forest Industry Group, Daxing’anling Forestry Group and Yichun Forest Industry Group as case studies to analyze the role of state-owned forest areas for otter conservation. </p><p id="p00015"><strong>Results:</strong> The results demonstrate that potential distribution and conservation priority areas cover 104,515.04 km<sup>2 </sup>and 45,448.99 km<sup>2</sup>, respectively. Large swathes of conservation priority areas remain for the otter in Daxing’anling which are connected with conservation priority areas in Xiaoxing’anling and there is no obvious geographical boundary between them. Therefore, these regions are mandatory to conserve to maintain the stability of otter populations in Northeast China. Otters in the Liaoning Province faced the greatest anthropogenic pressures, followed by the Jilin Province and Heilongjiang Province. Otters in the Inner Mongolia Autonomous Region faced the lowest amount of pressure. There is potential distribution for the otter in 63 nature reserves and conservation priority areas in 32 nature reserves out of the 110 reserves in the region. The study area includes 11.64% of the otter’s potential distribution and 10.88% of conservation priority areas. Three state-owned forest industry groups cover 71.18% of the potential distribution of the otter (74,390.89 km<sup>2</sup>) and 79.26% of the conservation priority areas (36,022.22 km<sup>2</sup>). </p><p id="p00020"><strong>Conclusions:</strong> This study indicates that state-owned forest areas may play a greater role in the protection of biodiversity following a comprehensive ban on logging in natural forests. Therefore, we propose that areas with important conservation value in state-owned forest land should be gradually included under the protections of national parks in order to achieve systematic and complete protection of biodiversity. Finally, we suggest: (1) strengthening the management of river pollutions; (2) reducing fishing intensity; (3) a long-term field monitoring network be established to study population changes in otters; (4) increasing the investment in scientific research on otters; and (5) raising awareness of otter conservation through public education.</p>
东北地区水獭分布格局与保护优先区识别
DOI:10.17520/biods.2021157
[本文引用: 3]
水獭是水生生态系统重要的指示种和旗舰种, 由于强烈的人为干扰, 中国的水獭种群数量大幅下降, 部分区域已局部灭绝。然而目前国内对水獭的调查和研究非常有限, 本底不清的状况已经严重影响到水獭的野外保育工作。本文以东北地区的欧亚水獭指名亚种(Lutra lutra lutra)为研究对象, 基于2016-2020年的调查数据, 使用组合建模的方法评估了水獭的潜在分布区; 利用地理信息系统和系统保护规划软件分析了水獭的保护优先区并计算了各省级行政区内水獭潜在分布区和保护优先区面临的人类压力; 结合国家级自然保护区的空间布局分析了水獭的保护现状, 并以内蒙古森工集团、大兴安岭林业集团、伊春森工集团三大国有林区为例分析了重点国有林区在水獭保护中的作用。结果表明: (1)水獭潜在分布区和保护优先区面积分别为104,515.04 km<sup>2</sup>和45,448.99 km<sup>2</sup>, 其中大兴安岭的水獭保护优先区集中连片, 并与小兴安岭的保护优先区相连, 栖息地之间没有明显地理隔离, 是维持东北地区水獭种群稳定的重中之重; (2)水獭面临的人类压力大小依次为: 辽宁 > 吉林 > 黑龙江 > 内蒙古; (3)研究区内110个国家级自然保护区中有63个包含水獭潜在分布区, 覆盖面积为12,168.93 km<sup>2</sup>, 仅占水獭潜在分布区面积的11.64%, 其中32个国家级自然保护区包含水獭保护优先区, 占水獭保护优先区面积的10.88%; (4)三大国有林区涵盖了71.18%的水獭潜在分布区和79.26%的保护优先区(面积分别为74,390.89 km<sup>2</sup>和36,022.22 km<sup>2</sup>)。由此可见, 尽管水獭潜在分布区中国家级自然保护区占比较低, 但是在天然林全面禁伐的背景下, 重点国有林区可能在未来东北地区的生物多样性保护中发挥更大作用, 因此我们建议将重点国有林区中具有重要保护价值的区域逐步纳入以国家公园为主体的自然保护地体系中, 以实现生物多样性的系统性和完整性保护。最后, 本文结合研究结果和实地调研提出以下保护建议: (1)加强对河流污染物的管理; (2)控制渔民捕鱼强度; (3)开展全面的水獭专项调查并建立长期的监测体系; (4)加大对水獭的科研投入; (5)加强宣传力度, 提升公众保护意识。
Biological invasions facilitate zoonotic disease emergences
DOI:10.1038/s41467-022-29378-2
PMID:35365665
[本文引用: 2]
Outbreaks of zoonotic diseases are accelerating at an unprecedented rate in the current era of globalization, with substantial impacts on the global economy, public health, and sustainability. Alien species invasions have been hypothesized to be important to zoonotic diseases by introducing both existing and novel pathogens to invaded ranges. However, few studies have evaluated the generality of alien species facilitating zoonoses across multiple host and parasite taxa worldwide. Here, we simultaneously quantify the role of 795 established alien hosts on the 10,473 zoonosis events across the globe since the 14 century. We observe an average of ~5.9 zoonoses per alien zoonotic host. After accounting for species-, disease-, and geographic-level sampling biases, spatial autocorrelation, and the lack of independence of zoonosis events, we find that the number of zoonosis events increase with the richness of alien zoonotic hosts, both across space and through time. We also detect positive associations between the number of zoonosis events per unit space and climate change, land-use change, biodiversity loss, human population density, and PubMed citations. These findings suggest that alien host introductions have likely contributed to zoonosis emergences throughout recent history and that minimizing future zoonotic host species introductions could have global health benefits.© 2022. The Author(s).
The neglected otters in China: Distribution change in the past 400 years and current conservation status
DOI:10.1016/j.biocon.2018.10.028 URL [本文引用: 1]
Historical distribution of the otter (Lutra lutra) in north-east China according to historical records (1950-2014)
DOI:10.1002/aqc.2624 URL [本文引用: 1]
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