
生物多样性 ›› 2025, Vol. 33 ›› Issue (11): 25203. DOI: 10.17520/biods.2025203 cstr: 32101.14.biods.2025203
沈君瀚1,2,3(
), 高海洋3(
), 孙松3(
), 吴飞3, 何南4, 王鹤5, 华彦3,*(
)(
), 武正军1,2,*(
)(
)
收稿日期:2025-05-29
接受日期:2025-09-21
出版日期:2025-11-20
发布日期:2025-12-26
通讯作者:
华彦,武正军
基金资助:
Junhan Shen1,2,3(
), Haiyang Gao3(
), Song Sun3(
), Fei Wu3, Nan He4, He Wang5, Yan Hua3,*(
)(
), Zhengjun Wu1,2,*(
)(
)
Received:2025-05-29
Accepted:2025-09-21
Online:2025-11-20
Published:2025-12-26
Contact:
Yan Hua, Zhengjun Wu
Supported by:摘要:
明确濒危物种的微生境选择特征是为其制定科学保护策略的基础。鳄蜥(Shinisaurus crocodilurus)是全球濒危物种, 野外种群数量仅剩1,500只左右。本研究于广东罗坑鳄蜥国家级自然保护区, 针对鳄蜥集中分布的溪沟开展微生境样方调查, 系统分析了鳄蜥的微生境选择特征。随机森林模型分析以及袋外误差计算表明栖枝密度、溪流深度、溪流沙覆盖比、郁闭度、溪流流速和溪流宽度是影响鳄蜥利用微生境的重要变量, 并具有特定的生态阈值。广义线性混合效应最终模型表明, 栖枝密度、溪流深度、郁闭度和溪流宽度与鳄蜥出现概率呈显著正相关; 其中, 栖枝密度为影响鳄蜥出现概率的首要因子, 溪流深度次之; 郁闭度虽独立贡献率较低, 然其作用不可或缺, 是基础性影响因子; 而溪流宽度则为边缘影响因子。多模型分析结果综合表明, 鳄蜥尤其偏好高栖枝密度的微生境, 可能与其获取庇护和食物资源的需求相关。研究建议, 在鳄蜥保护中应以提高栖枝密度构建多层次庇护网络作为首要目标, 同时通过改善水文环境和提高乔木郁闭度等措施, 为鳄蜥种群提供更适宜的生存环境。
沈君瀚, 高海洋, 孙松, 吴飞, 何南, 王鹤, 华彦, 武正军 (2025) 鳄蜥的微生境选择特征及保护启示. 生物多样性, 33, 25203. DOI: 10.17520/biods.2025203.
Junhan Shen, Haiyang Gao, Song Sun, Fei Wu, Nan He, He Wang, Yan Hua, Zhengjun Wu (2025) Microhabitat selection characteristics and conservation implications of the Chinese crocodile lizard (Shinisaurus crocodilurus). Biodiversity Science, 33, 25203. DOI: 10.17520/biods.2025203.
图1 广东罗坑鳄蜥国家级自然保护区鳄蜥种群生境及行为。(A)鳄蜥适宜微生境; (B)夜间鳄蜥静伏于栖枝睡眠; (C)日间鳄蜥静伏于岩石上晒太阳; (D)鳄蜥捕食蚯蚓。
Fig. 1 Habitat and behavior of the Chinese crocodile lizard (Shinisaurus crocodilurus) population in Guangdong Luokeng Crocodile Lizard National Nature Reserve. (A) Suitable microhabitat; (B) Nocturnal perching and resting on branches; (C) Basking on a rock during daytime; (D) Preying on an earthworm.
| 变量名 Variable | 变量介绍 Variable introduction | 测量方法 Measurement methods | 平均值 Mean | 中位数 Median | 标准差 SD | 频数 Frequency | |||
|---|---|---|---|---|---|---|---|---|---|
| 连续变量 Continuous variables | |||||||||
| 树冠层高度 Canopy height (m) | 从地面至乔木树冠顶部的垂直距离 Vertical height from the ground level to the top of the tree canopy | 采用2 m标准枝杆作为视觉参照标尺进行目测估算 Visual estimation was performed using a 2-meter standard branch pole as a visual reference scale | 8.93 | 8.10 | 3.65 | - | |||
| 灌木层高度 Shrub layer height (cm) | 从地面至灌木层顶部的垂直距离 Vertical height from the ground to the top of the shrub layer | 利用钢卷尺测量样方内5个随机选择的灌木斑块的高度, 并计算其平均值 The height of five randomly selected woody patches within the quadrat was measured using a measuring tape, and the average value was calculated | 84.70 | 77.00 | 36.70 | - | |||
| 郁闭度 Canopy closure (%) | 林冠层枝叶垂直投影面积占林地总面积的比例 Canopy cover percentage | 通过测量样方内4个角点及中心点的垂直冠层投影面积与样方总面积的比值, 并计算5个测量点的平均值来确定 It was determined by measuring the ratio of the vertical canopy projection area to the total quadrat area at the four corners and the center point within the quadrat, and then calculating the mean value from these five measurement points | 52.90 | 52.60 | 16.20 | - | |||
| 溪流深度 Stream depth (cm) | 溪流的平均深度 Mean depth of the stream | 在每个样方的上、中、下部测量, 包括最大/最小宽度和深度, 最终计算平均值 Measurements of maximum/minimum width and depth were taken at the upper, middle, and lower sections of each quadrat, with the mean values subsequently calculated | 19.30 | 17.10 | 9.77 | - | |||
| 溪流宽度 Stream width (cm) | 溪流的平均宽度 Mean width of the stream | 151.00 | 143.00 | 56.00 | - | ||||
| 溪流流速 Stream velocity (cm/s) | 溪流的平均流速 Mean flow velocity of the stream | 采用LS300-A型便携式流速仪测定样方上、中、下游的表面流速(cm/s), 测定值经算术平均与对数转换后用于统计分析 Surface flow velocity (cm/s) at the upstream, midstream, and downstream sections of the quadrat was measured using a portable LS300-A current meter. The measured values were arithmetically averaged and log-transformed for statistical analysis | 24.60 | 26.40 | 14.40 | - | |||
| 溪沟壁坡度 Gully bank slope (°) | 溪流两侧土坡或石壁的坡度 Streambank slope | 采用坡度仪测量样方上部、中部和下部的两侧溪沟壁坡度, 计算平均值 The gradient of both streambanks at the upper, middle, and lower sections of the quadrat was measured using a clinometer, and the mean value was calculated | 46.30 | 48.20 | 18.10 | - | |||
| 溪沟坡度 Gully slope (°) | 样方沿溪流的整体坡度 Overall longitudinal slope of the quadrat along the stream | 使用坡度计测量样方的4个角点和中心位置, 共5个点, 计算所得数据的平均值 The mean value was calculated from measurements taken with a clinometer at five locations: the four corners and the center of the quadrat | 8.72 | 7.67 | 6.42 | - | |||
| 溪沟坡向 Gully aspect (°) | 溪流流向的朝向 Flow direction azimuth | 使用罗盘(0-360°)进行测量。为处理统计分析中的循环特性, 将数据进行正弦变换处理 Measured using a compass (0-360°). To address the circular nature of the data in statistical analysis, the values were transformed using a sine transformation | -0.34 | -0.84 | 2.71 | - | |||
| 溪流沙覆盖比 Stream sand coverage percentage (%) | 溪流中沙在基质表面的覆盖比例 Percent sand cover on the streambed | 基于溪流宽度测量数据, 构建样地俯视平面图。将样方划分为0.5 m × 0.5 m的单元网格, 基于目视判读记录各网格属性参数, 通过网格法定量测算溪流沙质基底覆盖率、灌草盖度及溪流面积(关法春等, | 59.20 | 57.80 | 22.90 | - | |||
| 灌草盖度 Shrub and herb coverage (%) | 灌木层与草本层植被的垂直投影面积占样方内陆地总面积的百分比 Percent shrub and herb cover | 43.70 | 43.40 | 19.00 | - | ||||
| 溪流面积 Stream area (m²) | 溪流的面积 Area of stream | 6.67 | 6.20 | 2.94 | - | ||||
| 栖枝密度 Perch density (unit/m²) | 溪流水面上每平方米有多少根可供鳄蜥栖息使用的树枝 Density of branches suitable for Chinese crocodile lizard perching per square meter of stream water surface | 统计被认为是鳄蜥可用的悬垂于水面的栖枝数量(罗树毅等, | 0.86 | 0.71 | 0.90 | - | |||
| 逻辑变量 Logical variables | |||||||||
| 回水塘有无 Backwater pond existence | 有 Yes (1) | 样方里有没有小池塘 Presence of small ponds within the quadrat | 通过直接观察将其分类为1 (存在)或0 (不存在) It was categorized as 1 (present) or 0 (absent) through direct observation | - | - | - | 156 | ||
| 无 No (0) | - | - | - | 80 | |||||
| 字符变量 Character variables | |||||||||
| 林型 Forest type | 常绿阔叶林 Evergreen broad-leaved forest | 植被类型 Vegetation type | 依据实地评估, 将其归类为常绿阔叶林或竹林 It was classified as either evergreen broad-leaved forest or bamboo forest based on field assessment | - | - | - | 170 | ||
| 竹林 Bamboo forest | - | - | - | 66 | |||||
表1 广东罗坑鳄蜥国家级自然保护区鳄蜥种群微生境变量描述表
Table 1 Descriptive table of microhabitat variables for Chinese crocodile lizards in Guangdong Luokeng Crocodile Lizard National Nature Reserve
| 变量名 Variable | 变量介绍 Variable introduction | 测量方法 Measurement methods | 平均值 Mean | 中位数 Median | 标准差 SD | 频数 Frequency | |||
|---|---|---|---|---|---|---|---|---|---|
| 连续变量 Continuous variables | |||||||||
| 树冠层高度 Canopy height (m) | 从地面至乔木树冠顶部的垂直距离 Vertical height from the ground level to the top of the tree canopy | 采用2 m标准枝杆作为视觉参照标尺进行目测估算 Visual estimation was performed using a 2-meter standard branch pole as a visual reference scale | 8.93 | 8.10 | 3.65 | - | |||
| 灌木层高度 Shrub layer height (cm) | 从地面至灌木层顶部的垂直距离 Vertical height from the ground to the top of the shrub layer | 利用钢卷尺测量样方内5个随机选择的灌木斑块的高度, 并计算其平均值 The height of five randomly selected woody patches within the quadrat was measured using a measuring tape, and the average value was calculated | 84.70 | 77.00 | 36.70 | - | |||
| 郁闭度 Canopy closure (%) | 林冠层枝叶垂直投影面积占林地总面积的比例 Canopy cover percentage | 通过测量样方内4个角点及中心点的垂直冠层投影面积与样方总面积的比值, 并计算5个测量点的平均值来确定 It was determined by measuring the ratio of the vertical canopy projection area to the total quadrat area at the four corners and the center point within the quadrat, and then calculating the mean value from these five measurement points | 52.90 | 52.60 | 16.20 | - | |||
| 溪流深度 Stream depth (cm) | 溪流的平均深度 Mean depth of the stream | 在每个样方的上、中、下部测量, 包括最大/最小宽度和深度, 最终计算平均值 Measurements of maximum/minimum width and depth were taken at the upper, middle, and lower sections of each quadrat, with the mean values subsequently calculated | 19.30 | 17.10 | 9.77 | - | |||
| 溪流宽度 Stream width (cm) | 溪流的平均宽度 Mean width of the stream | 151.00 | 143.00 | 56.00 | - | ||||
| 溪流流速 Stream velocity (cm/s) | 溪流的平均流速 Mean flow velocity of the stream | 采用LS300-A型便携式流速仪测定样方上、中、下游的表面流速(cm/s), 测定值经算术平均与对数转换后用于统计分析 Surface flow velocity (cm/s) at the upstream, midstream, and downstream sections of the quadrat was measured using a portable LS300-A current meter. The measured values were arithmetically averaged and log-transformed for statistical analysis | 24.60 | 26.40 | 14.40 | - | |||
| 溪沟壁坡度 Gully bank slope (°) | 溪流两侧土坡或石壁的坡度 Streambank slope | 采用坡度仪测量样方上部、中部和下部的两侧溪沟壁坡度, 计算平均值 The gradient of both streambanks at the upper, middle, and lower sections of the quadrat was measured using a clinometer, and the mean value was calculated | 46.30 | 48.20 | 18.10 | - | |||
| 溪沟坡度 Gully slope (°) | 样方沿溪流的整体坡度 Overall longitudinal slope of the quadrat along the stream | 使用坡度计测量样方的4个角点和中心位置, 共5个点, 计算所得数据的平均值 The mean value was calculated from measurements taken with a clinometer at five locations: the four corners and the center of the quadrat | 8.72 | 7.67 | 6.42 | - | |||
| 溪沟坡向 Gully aspect (°) | 溪流流向的朝向 Flow direction azimuth | 使用罗盘(0-360°)进行测量。为处理统计分析中的循环特性, 将数据进行正弦变换处理 Measured using a compass (0-360°). To address the circular nature of the data in statistical analysis, the values were transformed using a sine transformation | -0.34 | -0.84 | 2.71 | - | |||
| 溪流沙覆盖比 Stream sand coverage percentage (%) | 溪流中沙在基质表面的覆盖比例 Percent sand cover on the streambed | 基于溪流宽度测量数据, 构建样地俯视平面图。将样方划分为0.5 m × 0.5 m的单元网格, 基于目视判读记录各网格属性参数, 通过网格法定量测算溪流沙质基底覆盖率、灌草盖度及溪流面积(关法春等, | 59.20 | 57.80 | 22.90 | - | |||
| 灌草盖度 Shrub and herb coverage (%) | 灌木层与草本层植被的垂直投影面积占样方内陆地总面积的百分比 Percent shrub and herb cover | 43.70 | 43.40 | 19.00 | - | ||||
| 溪流面积 Stream area (m²) | 溪流的面积 Area of stream | 6.67 | 6.20 | 2.94 | - | ||||
| 栖枝密度 Perch density (unit/m²) | 溪流水面上每平方米有多少根可供鳄蜥栖息使用的树枝 Density of branches suitable for Chinese crocodile lizard perching per square meter of stream water surface | 统计被认为是鳄蜥可用的悬垂于水面的栖枝数量(罗树毅等, | 0.86 | 0.71 | 0.90 | - | |||
| 逻辑变量 Logical variables | |||||||||
| 回水塘有无 Backwater pond existence | 有 Yes (1) | 样方里有没有小池塘 Presence of small ponds within the quadrat | 通过直接观察将其分类为1 (存在)或0 (不存在) It was categorized as 1 (present) or 0 (absent) through direct observation | - | - | - | 156 | ||
| 无 No (0) | - | - | - | 80 | |||||
| 字符变量 Character variables | |||||||||
| 林型 Forest type | 常绿阔叶林 Evergreen broad-leaved forest | 植被类型 Vegetation type | 依据实地评估, 将其归类为常绿阔叶林或竹林 It was classified as either evergreen broad-leaved forest or bamboo forest based on field assessment | - | - | - | 170 | ||
| 竹林 Bamboo forest | - | - | - | 66 | |||||
图3 广东罗坑鳄蜥国家级自然保护区鳄蜥种群关键变量筛选随机森林模型分析图。(A)随机森林模型微生境变量重要性分析图; (B-G) 6个关键变量部分依赖图。灰色区域表示无数据观测范围。
Fig. 3 Analysis of key variable selection for the Chinese crocodile lizard population in Guangdong Luokeng Crocodile Lizard National Nature Reserve. (A) Variable importance ranking from random forest model analysis for microhabitat selection; (B-G) Partial dependence plots for six key habitat variables. Grey areas indicate the range with no data observation.
| 变量名 Variable | 平均效应值 β_avg | 标准误 SE | Z | P |
|---|---|---|---|---|
| 截距 Intercept | -6.322 | 1.007 | -6.280 | 3.39e-10*** |
| 栖枝密度 Perch density (PD) | 1.906 | 0.315 | 6.052 | 1.43e-09*** |
| 溪流深度 Stream depth (SD) | 0.114 | 0.025 | 4.497 | 6.89e-06*** |
| 郁闭度 Canopy closure (CC) | 0.031 | 0.011 | 2.880 | 0.004** |
| 溪流宽度 Stream width (SW) | 0.008 | 0.004 | 2.080 | 0.038* |
表2 广东罗坑鳄蜥国家级自然保护区鳄蜥种群最优及最终广义线性混合模型统计描述表
Table 2 Optimal and final generalized linear mixed model statistical summary table for Chinese crocodile lizards in Guangdong Luokeng Crocodile Lizard National Nature Reserve
| 变量名 Variable | 平均效应值 β_avg | 标准误 SE | Z | P |
|---|---|---|---|---|
| 截距 Intercept | -6.322 | 1.007 | -6.280 | 3.39e-10*** |
| 栖枝密度 Perch density (PD) | 1.906 | 0.315 | 6.052 | 1.43e-09*** |
| 溪流深度 Stream depth (SD) | 0.114 | 0.025 | 4.497 | 6.89e-06*** |
| 郁闭度 Canopy closure (CC) | 0.031 | 0.011 | 2.880 | 0.004** |
| 溪流宽度 Stream width (SW) | 0.008 | 0.004 | 2.080 | 0.038* |
| 变量名 Variable | 平均效应值 β_avg | 标准误 SE | 置信区间下限 CI_Low | 置信区间上限 CI_High | 相对重要性 Relative importance (RI) | P |
|---|---|---|---|---|---|---|
| 截距 Intercept | -6.432 | 1.148 | -8.692 | -4.172 | < 0.001 | |
| 栖枝密度 Perch density (PD) | 1.825 | 0.324 | 1.190 | 2.460 | 1.000 | < 0.001 |
| 溪流深度 Stream depth (SD) | 0.120 | 0.027 | 0.068 | 0.172 | 1.000 | < 0.001 |
| 郁闭度 Canopy closure (CC) | 0.031 | 0.011 | 0.009 | 0.053 | 1.000 | 0.005 |
| 溪流宽度 Stream width (SW) | 0.008 | 0.004 | -0.001 | 0.016 | 0.747 | 0.051 |
| 溪流沙覆盖比 Stream sand coverage percentage (SSCP) | 0.011 | 0.008 | -0.005 | 0.027 | 0.506 | 0.160 |
| 溪流流速 Stream velocity (SV) | 0.144 | 0.161 | -0.172 | 0.459 | 0.315 | 0.372 |
表3 广东罗坑鳄蜥国家级自然保护区鳄蜥种群广义线性混合效应全局模型95%累计权重模型平均统计描述表
Table 3 Summary statistics of the 95% cumulative weight model averaging results from the global generalized linear mixed model (GLMM) for Chinese crocodile lizards population in Guangdong Luokeng Crocodile Lizard National Nature Reserve
| 变量名 Variable | 平均效应值 β_avg | 标准误 SE | 置信区间下限 CI_Low | 置信区间上限 CI_High | 相对重要性 Relative importance (RI) | P |
|---|---|---|---|---|---|---|
| 截距 Intercept | -6.432 | 1.148 | -8.692 | -4.172 | < 0.001 | |
| 栖枝密度 Perch density (PD) | 1.825 | 0.324 | 1.190 | 2.460 | 1.000 | < 0.001 |
| 溪流深度 Stream depth (SD) | 0.120 | 0.027 | 0.068 | 0.172 | 1.000 | < 0.001 |
| 郁闭度 Canopy closure (CC) | 0.031 | 0.011 | 0.009 | 0.053 | 1.000 | 0.005 |
| 溪流宽度 Stream width (SW) | 0.008 | 0.004 | -0.001 | 0.016 | 0.747 | 0.051 |
| 溪流沙覆盖比 Stream sand coverage percentage (SSCP) | 0.011 | 0.008 | -0.005 | 0.027 | 0.506 | 0.160 |
| 溪流流速 Stream velocity (SV) | 0.144 | 0.161 | -0.172 | 0.459 | 0.315 | 0.372 |
图4 广东罗坑鳄蜥国家级自然保护区鳄蜥种群最终模型分析图
Fig. 4 Final generalized linear mixed model analysis for the Chinese crocodile lizards population in Guangdong Luokeng Crocodile Lizard National Nature Reserve
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